Effects of two spring prescribed fires on epigeal Coleoptera in dry sclerophyll eucalypt forest in Victoria, Australia

Fores: ylogy Management Forest EcologyandManagement76 (1995) 69-85

Effects of two spring prescribed fires on epigeal Coleoptera in dry sclerophyll eucalypt forest in Victoria, Australia Nick G. Collett *, Fred G. Neumann Department

of Conservation

and Natural

Resources,

378 Cotham

Road

Kew, Vie. 3101, Australia

Accepted10 March 1995

Abstract The cumulative effects of two low-intensity prescribed fires, applied within 3 years during spring (15 October 1985 and 3 November 19881, were assessed on families and species of surface-active Coleoptera (beetles) in litter of dry sclerophyll mixed eucalypt forest over a 5.7 year period between 26 March 1985 and 26 November 1990. The study was based on 13 925 adult specimens, representing 31 families and 105 species, of which 70 species (66.7% of total) are undescribed. The most commonly trapped ‘major’ families were the predominantly predatory Staphylinidae and the decomposer/fungus feeding Nitidulidae and Leiodidae. The specimens were contained in 2240 pitfall trap samples collected from a burnt and an unburnt (‘control’) site adjacent to each other within a 30.7 ha study area. The two fires had no discernible cumulative effect on the activity of the Nitidulidae, the Leiodidae, the pool of 28 rarely trapped ‘minor’ families and the totals of decomposers/fungus feeders. However, short bursts in activity occurred among the first three of these taxa/groups during periods between the two fires. l.n contrast, activity of the Staphylinidae, the most abundant of all taxa trapped, declined significantly after the second fire relative to pre-fit fire levels. This trend accounted for the observed reductions in activity of total Coleoptera, of total predators and of the proportions of Coleoptera in total arthropods. Family and species richness of the Coleoptera remained unchanged following the fires. An undescribed fungus-feeding lathridiid (Aridius sp.) incurred a short upsurge in activity after the second fire. As only activity (and not diversity and taxon richness) of the Staphylinidae was affected by the fires, it is expected that they will return to pre-fire activity levels when the forest recovers in the absence of further fires. Two short-rotation low-intensity spring fines can therefore be applied in the forest type studied without causing unacceptable adverse impacts on litter-frequenting Coleoptera. Keyworak

Staphylinidae;Nitidulidae;Leiodidae;Activity; ‘Burnt’ site; ‘Control’ site

1. Introduction Repeated low-intensity prescribed fire applied during cool weather at strategic localities in Victoria’s

* Correspondingauthor.

3.7 million ha of dry sclerophyll mixed species eucalypt forest is considered to be an effective and economic fire protection measure. It reduces highly flammable fine fuels (humus, leaves, bark, desiccated ground vegetation and twigs of less than 6 mm diameter) (Luke and McArthur, 1978) on the forest floor, and provides low hazard barriers that reduce the frequency, rate of spread and intensity of wild-

0378-1127/95/$09.50 0 1995Elsevier ScienceB.V. All rights reserved SSDI 0378-1127(95)03560-5

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/Forest

fires and their associated damage to forest ecosystems (Johnston et al., 1983; Rawson et al., 1983; Department of Conservation, Forests and Lands, 1986; Greig, 1986). In Victoria, prescribed fire operations commenced during the 195Os, and an average of just under 130000 ha of State forest, national parks and other protected land have been burnt annually since 1983 (Forests Commission Victoria, 1983; Department of Conservation, Forests and Lands, 1983/89; Department of Conservation and Environment, 1989/91). However, there is considerable concern that such practice induces adverse side-effects on the physical, biotic and aesthetic attributes of forest ecosystems-an important issue that forest managers must address and, if necessary, ameliorate (Pratten, 1985; New, 1987; Christensen and Abbott, 1989). The Coleoptera represent a substantial part of Australia’s insect fauna with over 20000 species described. They are a major and ecologically important component of the epigeal arthropod fauna in the litter of dry sclerophyll forest (Collett et al., 1983; Neumann and Tolhurst, 19911, representing a wide range of feeding types including herbivores, fungus and seed feeders, predators, and decomposers (Booth et al., 1990; Lawrence and Britton, 1991). Regarding the concern by Springett (1976) and Majer (1984) that prescribed fire may decimate surface-active Coleoptera and thus adversely affect litter decomposition, nutrient cycling and biological control, Neumann and Tolhurst (1991) found that a single prescribed low-intensity fire in dry sclerophyll forest during spring or autumn had no discernible positive or negative effects on total Coleoptera at litter level. Subsequently, Collett et al. (1993) found in the same forest that two prescribed fires within 3 years in spring had progressively lowered coleopteran activity after each fire, a trend which culminated in a significant decline (P < 0.05) being recorded between the pre-first fire and the post-second fire activity levels. As a simultaneous drop in activity was also detected at the unburnt (‘control’) site, it was unclear whether (1) cumulative fire effects alone had caused the decline, and (2) the decline involved a single or a range of coleopteran taxa. The present paper addresses these uncertainties by examining the Coleoptera at family and species level relative to the cumulative effects of the two prescribed fires.

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76 11995) 694.5

2. Methods 2.1. Study sites Two adjacent forest sites, designated as ‘control’ (13.5 ha) and ‘burnt’ (17.2 ha) were selected for study in a dry sclerophyll eucalypt forest with an average annual rainfall of 905 mm. The study area is known locally as ‘Blakeville’ (375l’ S, 144”lO’ E) and is located approximately 18 km south of the city of Daylesford in south-central Victoria. The ‘burnt’ site was fire-treated on 15 October 1985 (thereafter called the ‘first fire’) and again on 3 November 1988 (the ‘second fire’), whereas the ‘control’ site remained unburnt. The fires were lit during mid-moming periods using a 50 m X 50 m grid of spot ignitions The topography at both sites is gently undulating, the aspect is predominantly south-westerly, and the elevation of the forested landscape is 625-700 m above sea level. At the beginning of sampling the Coleoptera in 1985, both sites had not been burnt since 1935, and the mean depth of surface fuel (humus, litter and twigs of < 26 mm in diameter) on the forest floor was around 35.8 mm (n = 1001. The projected foliage cover of the overstorey and understorey was approximately 66% and 35% respectively, and the vegetation at both sites was similar with a total of 110 plant species being recorded (Neumann and Tolhurst, 1991). No logging had occurred since 1935, but some thinning to remove trees of low commercial value had been carried out during 1961 and 1964. In 1985 the basal area of the overstorey was 25.8 m2 ha -’ and of the regrowth 7.8 m’ ha -! (Tolhurst and Flinn, 1992). The overstorey, 35 m tall, was dominated by Messmate Stringybark (Eucalyptus obliqua L’Herit), Narrow-leaf Peppermint (EucaZypus rudiutu Sieber ex DC), and Candlebark (Eucalypbus rubida M. Deane&Maiden), whereas the understorey was generally less than 2 m tall and composed mainly of Silver Wattle (Acacia dealbata Link.), Hop Wattle (Acacia stricta (Andr.) Willd.), Austral Bracken (Pteridium esculentwm (Forst. F.) Cockayne), Forest Wire-grass (Tetrurrhena juncea R. Br.), Tussock Grass (Pea sieberiana Sprengel), wallaby grasses (Danthonia spp.), Common Raspwort (Gonocurpus tetragyws Labill.), and Ivy-leaf Violet (Viola h&zh-

N.G. Collett,

F.G. Neumann/Forest

acea Labill.). The nomenclature of these species was based on Forbes and Ross (1988). 2.2. Sampling The prevailing weather conditions and the monthly soil dryness index (SDI) values were recorded from 1 January 1986 to 31 December 1990 at a computerized weather station (Model EASIDATA MK-II; Environdata Aust.) in an open unburnt area within ‘Blakeville’ forest approximately 3 km from the study sites. SD1 values indicate the amount of rain required to bring the soil profile back to field capacity expressed on a O-200 scale, where 0 is field capacity and 200 is worst possible drought (Mount, 1972). The forward rate of spread of the fires was estimated at each site by recording the time taken by the fire front to move between 16 posts in a line at 2 m apart within a 30 m X 35 m permanent plot, and by measuring the distance between numbered metal plates placed along the fire front at known times in various accessible locations. Scorch height and the proportion (%b) of area burnt were assessed within a week of each fire at 20 sampling points on a 50 m X 100 m grid by measuring the highest scorch mark on a tree within a 20 m radius of each sampling point, and by placing a 10 m tape on the ground and measuring in terms of 10 cm long units, the length of tape that intercepted burnt patches respectively. Fire line intensity (I) (kW m-l > was calculated using the formula I = whr, where w is weight of fuel burnt (kg), h is calorific value of fuel (18600 kJ kg-‘) and r is forward rate of spread of fire (m s-l> (Byram, 1959). Surface soil temperatures during the fires were estimated at the burnt site within a 0.1 ha plot by means of 12 0.3 mm thick aluminium plates, marked with heat sensitive Thermochrom (Faber-Castell) crayons. Each plate was folded into a 12 cm X 5 cm envelope and placed between the litter base and the soil surface. The crayons were sensitive to temperatures between 65°C and 600°C. Fine fuels were sampled from this plot 1 month before burning, within a week after burning, and then annually for 3 years during spring. The unburnt ‘control’ site was similarly sampled. On each sampling occasion, two replicates of 15 0.1 m3 fine fuel samples were

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collected from the ‘burnt’ site and one replicate of 15 samples from the ‘control’ site. The fuel samples were oven-dried at 105°C for 24 h to ensure constant weight among the samples. Adult Coleoptera were sampled contemporaneously at both study sites within a 1 ha permanent plot over 56 weekly periods between 26 March 1985 and 26 November 1990, i.e. over 5.7 years. Through the centre of each plot, 20 pitfall traps were positioned at 5 m intervals along a fixed 100 m transect. There were 2240 samples (20 pitfall traps X 56 weekly sampling periods X 2 sites). The forest at both sampling sites was similar in age, aspect, elevation vegetation and wildfire/ logging history. Each trap consisted of an 18 mm diameter test tube in a PVC sleeve. In previous studies by F.G.N. this diameter proved to be effective in preventing overflow of the traps during rainstorms, yet it did not appear to exclude the larger Coleoptera. The traps were half filled with 75% methanol and each provided a 2.5 cm2 receptive surface area. Digging-in effects were minimised by commencing sampling 2 weeks after positioning of the traps. Coleopteran activity was measured as the number of individuals trapped per composite sample of 20 pitfall traps over a 7 day period at the litter surface. Pitfall trapping estimates the relative population levels of surface-active invertebrates, and thereby measures their relative importance on the forest floor (Greenslade, 1964; Greenslade and Greenslade, 1971). The technique, when used with methanol preservative, is especially suitable for trapping adult beetles (Greenslade and Greenslade, 1971). Pitfall trapping was therefore considered appropriate for this study, as relative sampling over time was used to assess the effects of two low-intensity prescribed fires on the activity of litter-frequenting adult Coleoptera. 2.3. Analysis The trapped beetle specimens were counted and classified to family and species level. All other arthropod specimens trapped were also counted to provide examples of the proportions of the Coleoptera among total arthropods at litter level in twice burnt and unburnt forest. The nomenclature of coleopteran families was based on that given by Lawrence and

72

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Britton (19911, except that the scolytids were assigned family status in accordance with common practice in Europe and North America where these insects are abundantly represented. Booth et al. (1990) and Lawrence and Britton (1991) were consulted to identify the coleopteran families by feeding type. The staphylinids were considered to be predacious, although some of them feed on algae and fungi. Because the range of species and their activity varied substantially between the 20 trap locations at each site on many sampling occasions, each set of 20 contemporaneous trap collections was pooled into a composite sample. For the ‘control’ site, the mean annual activity between Autumn 1985 and Summer 1989/90 of total adult Coleoptera, of ‘major’ families and of the pool of ‘minor’ families was calculated from the 8-11 available composite samples per year. Any differences between years were tested by analysis of variance (ANOVA) after log,,(x + 1) transformations of the raw data to account for zeros in the data base, followed by the Tukey multiple comparison test for unequal sample sizes (Zar, 1984). Differences in seasonal activity of total Coleoptera and of the ‘major’ and ‘minor’ families between Autumn 1985 and Spring 1990 were similarly analysed. The effects of fire on the Coleoptera were assessed independently for the burnt site relative to the ‘control’ site by using a BACI design (Stewart-Oaten et al., 1986), which is an appropriate way to treat data of a ‘two sites (control/impact) and several times’ study (Green, 1993). In the present study, the set of differences between the contemporaneous observations over 7 day periods in the fire treatment and control series before burning were compared with those after burning for various parameters, using the Mann-Whitney non-parametric U-test (Zar, 1984). The use of differences eliminated the effects of site factors and seasonality and strengthened the validity of the assumption in the Mann-Whitney test that consecutive observations are independent. Parameters examined in the comparisons were: (1) the proportions of Coleoptera in composite samples of total arthropods at the litter surface of the ‘control’ and the ‘burnt’ site; (2) the Shannon-Wiener diversity index (Poole, 1974), the Margalef taxon richness index (Southwood, 1978), and Pielou’s community

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evenness~index (Pielou, 19661 for beetle families; (3) the activity at the litter surface of total Coleoptera, of commonly trapped ‘major’s families, of the pool of the rarely trapped ‘minor’ families, of total predators and of total decomposers/fungus feeders. Although the three ecological indices tested have certain limitations (Hurlbert, 1971), they were considered appropriate here because sampling was relative over time, restricted to sites in close proximity to each other and within the same forest type, and not aimed at a complete census of the coleopteran community. For each parameter, regression analyses were done on the control/impact differences against time to test for constancy among them as confirmation or rejection of fire impacts (Stewart-Oaten et al.. 1986). To determine the fire effects on species richness of the Coleoptera, two-dimensional chi-square analyses were performed on the species totals recorded before and after each fire at the ‘burnt’ and ‘control’ site (2 x 2 contingency tables). Also, three-dimensional chi-square tests (4 X 2 X 2 contingency tables) were carried out to test for mutual independence between the variables: (1) species frequency within selected coleopteran families/groups, (2) study sites, (3) fire treatments (Zar, 1984).

3. Resuits 3.1. Weather conditions

The mean monthly maximum air temperature at 1.5 m above the ground in a standard screen varied from 7.6”C in mid-winter (July 1987) to 25.7”C in mid-summer (January 19881, and the mean minima from 0.4”C (July 1987) to 10,3”C (February 19901. Total annual rainfall recorded in a 203 mm diameter collector at 1 m above the ground was 817 mm in 1986, 950 mm in 1987, 857 mm in 1988 998 mm in 1989 and 868 mm in 1990; total monthly rainfall ranged from a minimum of 2.5 mm in mid-summer (January 1990) to a maximum 173 mm in late summer (February 1990, associated with thunderstorm activity). However, the driest period (mean rainfall of 33.2 mm> occurred between late summer and early autumn (February, March) and the wettest (mean 101.6 mm) between late autumn~and midwinter (May-July). Monthly soil temperatures at

N.G. Collett,

F.G. Neumann/Forest

Ecology

l-2 cm depth below a 50 mm grass cover, and monthly soil dryness indices, are shown in Fig. 1. Comparative soil temperature data for the forest are not available, but some measurements by Tolhurst and Turvey (1992) have indicated that temperatures in the forest soil are likely to vary over a narrower range than that shown in Fig. l(a). Average weather conditions at the time of each experimental fire were as follows: air temperature, 18.W (‘first fire’> and 16.O”C (‘second fire’); relative humidity, 65% (‘first fire’) and 55% (‘second fire’); wind speed at 10 m above the forest floor,

and Management

76 (199.5) 69-85

73

10.5 km hh’ (‘first fire’) and 9.0 km h-l (‘second fire’); SD1 values, 16 (‘first fire’) and 37 (‘second fire’) (Tolhurst and Flinn, 1992). 3.2. Fire and fuel load statistics

The main difference between the first and second fires was their effect on surface soil temperature (Table 1). It was far higher during the ‘second fire’ probably because the litter was then relatively fresh, less compacted and without a thick decomposed layer of low flammability, thus allowing greater heat

(a) 30 25-

1

1986

+

--+--

1987

1988

__.. 0 ___.

--iI---

1989

01

1 Jan Feb

Mar

Apr May

Jun

Jul

Aug Sep

Ott

Nov

Dee

(b)

2x 2 .a

loo

+

1986

-4----,

1987

_-_- 0 _---

,988

--A----

1989

--a---

1990

.Z 5: .

50

Jan Feb Mar Apr May Jun Jul Aug Sep Ott Nov Dee Fig. 1. Monthly soil temperatures (a) and monthly soil dryness indices (b) recorded in an open grass-covered the study sites in west-central Victoria between 1 January 1986 and 31 December 1990.

area, approximately

3 lun from

74

N.G. Collett, F.G. Neumann/Forest Ecology and Managemenf 76 (199.5) 69-X’;

t ha-’ which is the acceptable level for fire prevention in Victoria, as stipulated by the Department of Conservation and Natural Resources.

Table 1 Data on fire behaviour for the two experimental spring fires applied in the study area on 15 October 1985 and 3 November 1988 Parameter Fire line intensity (kW m-l) Forward rate of spread (m min- ’ ) Surface soil temperature (“a Scorch height h-d Area burnt (%)

First fire

Second fire

Mean Max.

209 (47) 647 0.83 (0.12)

137 (64) 692 0.58 (0.14)

Mean Max. Mean Max.

65 (10) 100 6.5 (0.8) 17.0 74

400 (15) 500 4.5 (0.5) 12.5 86

Source: Collett et al. (1993). Values in parentheses refer to standard

3.3. Oueruiew of coleopteran families and species trapped

errors of means.

transfer to the soil (K. Tolhurst, personal communication, 1993). The ‘second fire’ was slower moving than the first, thus explaining why the fire line intensity and the scorch height of the first fire exceeded those of the second (Table 1). The fine fuel load on the ‘burnt’ site just prior to the ‘first fire’ was 16.2 (standard error 1.1) t ha-’ compared to 14.7 (0.7) t ha-’ before the ‘second fire’, whereas the immediate post-fire loads were 6.9 (0.6) t ha-’ and 5.7 (0.6) t ha-’ respectively (Fig. 2). Approximately similar amounts of fuel were therefore burnt during both fires. Moreover, both spring fires reduced the fine fuel loads below the threshold of 12.0

20

1

A total of 31 families and 105 species were identified among the 13925 specimens of adult Coleoptera trapped; 26 families and 79 specieswere recorded at the ‘control’ site and 25 families and 78 speciesat the ‘burnt’ site (Table 2). Seventy of the total species (66.7%) are still undescribed, and 12 (11.4%) could not be identified below sub-family rank (Table 3). Twenty families (64.5% of total) were common to both sites.Among the Staphylinidae, Nitidulidae and Leiodidae, the number of total specimenstrapped ranged from 970 to 6950 (Table 2). These families were therefore referred to as ‘major’ families to distinguish them from the 28 lesscommonly trapped ‘minor’ families. Six families were exclusive to the ‘control’ site (Ptiliidae, Throscidae, Coccinellidae, Mycetophagidae, Colydiidae, Aderidae? and five families to the ‘burnt’ site (Eucnemidae, Lycidae, Melyridae, Zopheridae, Mordellidae). Of these II families, fewer than four specimensper family- were trapped, indicating that they were either uncommon components of the surface active coleopteran fauna

Fire

Fire

t

0' 0

1

2

Time Fig. 2. Levels site.

3

4

5

(years)

of fine fuels on the forest floor before and after the two fires in spring at the ‘burnt’

site compared

with the nearby

‘CQHX&

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F.G. Neumann

/Forest

in the study forest, or that pitfall trapping was an unsuitable technique for collecting these families. The majority of the Coleoptera were decomposers, fungus feeders and predators, but a few were bark and wood borers, scavengers, omnivores and herbivores (Table 2). Fifty-two species (49.5% of total) were found at both study sites, and 93 (88.5%) belonged to the 20

Table 2 Statistics of the Coleoptera

trapped

at the ‘burnt’

and ‘control’

Family

Feeding

Staphylinidae a Nitidulidae Leiodidae Lathridiidae b Carabidae Cryptophagidae Curculionidae Anthicidae Bostrichidae Silvanidae Scarabaeidae Scydmaenidae Clambidae Histeridae Pselaphidae Tenebrionidae Chrysomelidae Anobiidae ’ Scolytidae Elateridae MeIyridae Mycetophagidae Ptiliidae Eucnemidae Throscidae Lycidae Coccinellidae Colydiidae Zopheridae Mordellidae Aderidae

Predators, fungus feeders Decomposers, fungus feeders Decomposers, fungus feeders Fungus feeders Predators Fungus feeders Herbivores, wood borers Decomposers (scavengers) Wood borers (sapwood only) Decomposers, fungus feeders Herbivores Predators Fungus feeders Predators Predators Decomposers (scavengers) Herbivores Bark/wood borers Bark borers Decomposers, predators Omnivores Fungus feeders Fungus feeders Decomposers, fungus feeders Fungus feeders Decomposers Predators Fungus feeders Fungus feeders Decomposers, herbivores Herbivores

Total beetles Total families Total species a b ’ d

type d

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76 (1995)

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families that were trapped at both sites. Species richness was found to be greatest among the Curculionidae, Staphylinidae, Scarabaeidae, Leiodidae and Chrysomelidae with eight or more species trapped per family and least among the Histeridae, Ptiliidae, Clambidae, Eucnemidae, Throscidae, Lycidae, Melyridae, Silvanidae, Cryptophagidae, Coccinellidae, Colydiidae, Zopheridae, Mordellidae and

study sites over a 5.7 year period from 26 March No. specimens and species (parentheses) from study site ‘Control’

‘Burnt’

2987 (10) 1688 (5) 455 (7) l(l) 66 (7) 121(l)

3963 3356 515 204 93 37 114 23

38 03) 36 (3) 11(2) 8 (1) 8 (6)

8 (2) 9 (1) 6 (1) 3 (3) 6 (4) 3 (4) 4 (2) 3 (2) 4 (2) 0 (0) 2 (2) 0 (0) l(1) l(l) 0 (0) l(1) l(1) 0 (0) 0 (0) l(l) 5472 26 79

Includes Scaphidiidae (Lawrence and Britton, 1991). 200 collected at the ‘burnt’ site between November and December 1988. Includes Ptinidae (Lawrence and Britton, 1991). Based on data given in Lawrence and Britton (1991) and Booth et al. (1990).

Totals

(11) (5) (7) (4) (6) (1) (9) (3)

46 (2) 310)

16 (6) 9 (1) 4 (1) 5 (1) 8 (3) 4 (3) 7 (4) 4 (2) 5 (1) 2 (2) 3 (1) 0 (0) 10) 0 (0) 0 (0) 10) 0 (0) 0 (0) 10) l(l) 0 (0) 8453 25 78

1985 to 26 November

6950 (11) so44 (5) 970 (8) 205 (4) 159 (7) 158 (1) 152 (14) 59 (3) 57 (2) 39 (1) 24 (9) 17 (2) 13 (1) 11(l) 11(3) 10 (5) 10 (8) 8 (2) 8 (2) 6 (4) 3 (1) 2 (2) l(l) l(1) l(l) l(1) l(1) l(1) l(1) l(1) l(l) 13925 31 105

1990

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Table 3 Species of Coieoptera by family trapped from 26 March 198.5 to 26 November 15 October 1985 and 3 November 19881 and of the ‘control’ site Family a or subfamily

Genus/species

‘Burnt’

CARABIDAE Broscini Psydrini Pterostichini

Eurylychnus blagrauei (Castelnau) Mecyclothoraxpunctipernnis (Macleayf Notonomus chalybeus (Dejean) Prosopognlus sp. Simodontus sp. 1 Simodontus sp. 2 Simodontus sp. 3

HISTERIDAE

Teretrius

PTILIIDAE

Genus indeterminate

LEIODIDAE Leiodinae

Pm-first

76 (1995)

69-85

1990 at the litter surface of the ‘burnt’

fire

Between ‘Control’

x

fires

study site (burnt

Post-second

fire

‘Burnt’

‘Control’

‘Burnt’

‘Control

X

x

X

\I

X

x

x

x

sp.

X

x

X

Y

x

x

‘r

x

I(

x

X

x

X

x

x

x

?.

X

Y

Y

b Colenisia sp. Zeadolphus sp. Colon sp. Austronemadus sp. Catoposchema sp. Nargomorphus globulus Nargomorphus sp. 1 Pseudonemadus sp.

Coloninae Cholevinae

X X

(Jeanne11

k

x

X

X

X

X

R

X

X

X

X

x

X

X

X

X

X

X

x

X

x x

X

X

X

X

SCYDMAENIDAE! Euconnus Euconnus STAPHYLINIDAE Oxytelinae

Osoriinae Paederinae Tachyporinae Staphyllininae Aleocharinae

sp. 1 sp. 2

X

x

b

Anotylus sp. 1 Anotylus sp. 2 Anotylus sp. 3 Osorius cktoriae (Oke) Hyperomma sp. Sepedophilus sp. Quedis sp. Aleochara sp. Atheta sp. Ocalea sp. Genus indeterminate

x

X

X

X

X

x

X

X

X

x

X

X

X

X

x

Anabaxis sp. Palimbolus sp. Pselaphaulax sp.

X

X

X

x

r;

x X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

x x x

X

x

x

X

X

X

X

X

X

X

x

s

PSELAPHIDAE

CLAMBIDAE SCARABAEIDAE Melolonthinae

Clambus

myrmecophilus

X

(Lea)

Heteronyx laeuiceps (Blackbum) Heteronyx terrenus (Blackbum) Heteronyx sp. Liparetrus erythropterus (Blanchard) Telura petiolata (B&on) Telura vitticollis (Erichson) Scitaia sericans (Erichson)

X

x

x x x Y

X

X

x

X

X

X

X

X X X

X X X x

on

X X

s

’

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Table 3 (continued) Pre-first

Between

Post-second

Genus/species

Scarabaeinae Dynastinae

Onthophagus Adoryphoras

longipes (Paulian) couloni (Burmeisterl

EUCNEMIDAE

Euryptychus

sp.

THROSCIDAE

Aulonothrosus

ELATERIDAE

Agrypnus caliginosus (Boisduval) Agrypnus sp. 1 Conoderus cordieri (Le Guilluo) Hapatesus electus (Neboiss)

x

Trichalus

x

‘Burnt’

LYCIDAE

fire

fires

Family a or subfamily

‘Control’

‘Burnt’

‘Control

‘Burnt’

fire ‘ Control’ x

x x

sp.

x

sp.

BOSTRICHIDAE Xylion Xylion ANOBIIDAE

MELYRIDAE NITIDULIDAE

SILVANIDAE

collaris (Erichson) lindi (Blackbum)

Dicranolaius Brachypeplus Carpophilus Thalycrodes Thalycrodes Thalycrodes

sp.

CRYPTOPHAGIDAECryptophagus’ COCCINELLIDAE

Diomus

(Lea)

bellulus (Guerin-Meneville) sp. sp. au&ale (Gemarl cylindricum (Blackbum) pulchrum (Blackbum)

Cryptamorpha

x x

x

x

X

x

x

Deltocryptus sp. Ptinus cupreoniger

b

x x

X X

x

x x x

x x

x

sp.

X

x

x

x

x

x

X

x x x

x x x

X

X

x

X

X

X

x

x

X

X

x

x

x

X

x

sp.

LATHRIDIIDAE Aridius sp. Cartodere constricta Corticaria sp. 1 Corticaria sp. 2

X

x x x

(Gyllenhal)

X

x

x

MYCETOPHAGIDAE Lifargus Typhaea

sp. stercorea

MORDELLIDAE

Mordellistena

COLYDIIDAE

Pycnomerus

ZOPHERIDAE

Cotulades

TBNEBRIONIDAE Helaeini Adeliini

Alleculini ANTHICIDAE

x

(Linnaeus)

X

sp.

X

sp.

x

sp.

x

Lepispilus rotundicollis (Blackbum) Adelium sp. 1 Adelium sp. 2 Seirotrana sp. Alcmeonis rufovittis (Carter) Anthicus sp. Pseudotomoderus Trichananca sp.

x

X

x X X

X X

X

sp.

x

X X

x

X

X

X

X

78

N.G. Collett,

F.G. Neumann/Forest

Ecology

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76 (1995) 69-X.5

Table 3 (continued) Family a or subfamily

Genus/species

ADERIDAE

Syzeton

CHRYSOMELIDAJZ Cryptocephalinae Eumolpinae Chrysomelinae

Ahicinae

SCOLYTIDAE!

CURCULIOMDAE Rhythirrinae Rhytirhininae Amycterinae Cryptorhynchinae

Entiminae

Total families Total species

Pre-first ‘Burnt’

fire

Between ‘Control’

fires

‘Burnt’

sp.

‘Burnt’

-

fire ‘Control’

x

Aporocera sp. Eboo sp. Edusella uiridicollis (Lcfevre) Paropsisterna? morio (Fabricus) Calomela bartoni (Baly) Calomela maculicollis (Boisduval) Arsipoda variegata (Waterhouse) Longitarsus uictoriensis (Blackburn) Xyleborus sp. Hylastes ater (Paykull)

Genus indeterminate Desiantha sp. Acantholophus sp. Genus indeterminate Genus indeterminate Genus indeterminate Genus indeterminate Genus indeterminate Genus indeterminate Genus indeterminate Genus indeterminate New genus Mandalotus sp. 1 Mandalotus sp. 3

‘Control’

Post-second

x

x x

’

x x sp. sp. sp. sp. sp. sp. sp. sp.

1 2 3 4 5 6 7 8

x x x

s

x

x x x

x x x x x

x

x Y

x

11 21

10 24

20 54

22 50

x x

Y

24 60

19 Sh

-.

a The families are in sequence of their phylogenetic affinity (Lawrence and B&ton, 1991). b These families are considered ‘major’ families wheras all others are termed ‘minor’ families. ’ This species is an introduced bark beetle specific to conifer forests (Neumann, 1987).

Aderidae, with only one species trapped per family (Table 2). At the ‘control’ site, the means of annual activity of total Coleoptera, of the ‘major’ families (Staphylinidae, Nitidulidae, Leiodidae) and of the pool of the 23 ‘minor’ families, were similar for the 5 years between 26 March 1985 and 26 February 1990 (F values for the log,,(x + 1) transformations of the data sets in each column of Table 4(a) were insignificant throughout). In contrast, some of the means of seasonal activity among total Coleoptera, the ‘major’ families and the pool of ‘minor’ families differed significantly (Table 4(b)). Summer was the least favoured season for activity of the three ‘major’

families, whereas it was the most active (together with spring and autumn) for the pool of the 28 ‘minor’ families. Low activity of the majority of Coleoptera during summer coincided with high average soil temperatures and moderately high soil dryness indices in late summer (Figs. l(a) and l(b)); 3.4. Effects of the fires The proportions of adult beetles in camposite samples of total surface-active arthropods trapped between 26 March 1985 and the time of the ‘first fire’ were similar to those observed over the period between the two fires at the ‘burnt’ site relative to

N.G. Collett,

F.G. Neumann

/Forest

Ecology

Year (MarchFebruary) or season (a) Annual 1985-1986 1986-1987 1987-1988 1988-1989 1989-1990

No. composite samples

actiuiq

a 11 10 8 11 9

(b) Seasonal actiuity ’ Autumn 12 Winter 13 Spring 12 Summer 12

Total Coleoptera

116.3 + 17.4 b

76 (1995)

and of the pool of ‘minor’ Mean no. adult specimens

families

per composite

Leiodidae

12.9 12.1

5.5 + 1.6

* 14.4 f 16.5

6.7 + 2.6 6.7 5 1.8

+ 10.1

4.1 + 1.4

5.6 + 1.3 7.4 + 2.2 10.2 + 2.2

+ 13.5 f 32.8

37.9 30.1

* 15.4 * 9.1

112.6 83.1

+ 33.0 $- 15.7

61.7 35.9

+ 18.9 + 7.3

71.4

f 14.4

29.3

f

38.5 33.1 27.8

76.4 33.8

f 17.2 a f 7.6 b

4.7

+ 10.4 ab 96.1 + 21.4 a 70.6 + 19.2 ab 18.8 f 5.2 b 44.5

‘Minor’ families

+ 2.7 + 3.3

60.8 90.3

13.3 + 13.3 +

4.1 b 2.2 b

‘control’

sample of

Nitidulidae

+ 38.5

i 21.7 a 142.5 + 26.8 a 102.5 + 18.1 ab 45.4+ 6.7 b

at the litter surface of the unburnt

Staphylinidae

136.1

137.3

79

69-85

The above results were consistent with the overall trend at the litter surface that pre- and post-fire activity at the ‘burnt’ site relative to the ‘control’ site was similar after each fire for total Coleoptera, the ‘major’ families Staphylinidae, Nitidulidae and Leiodidae, and the pool of the 28 ‘minor’ families, as well as for total predators and total decomposers/ fungus feeders (Tables 3 and 5). However, the activities of total Coleoptera, of the ‘major’ family Staphylinidae, and of total predators were significantly lower during the post-‘second fire’ period compared to pre-‘first fire’ levels (Table 5 and Figs. 3(a), 3(b) and 3(f)). In contrast, bursts of activity at the ‘burnt’ site far above those at the ‘control’ site, occurred among the ‘major’ families Leiodidae and Nitidulidae during the first winter and second spring following the initial fire respectively (Figs. 3(c) and 3(d)). The pool of the 28 ‘minor’ families also incurred an immediate post-‘first fire’ upsurge in activity which lasted until early summer 1986 (Fig. 3(e)), while the decomposers experienced an upsurge during the second spring following the ‘first fire’ (Fig. 3(g)). After the ‘second fire’, no short-term boosts in activity were observed among any taxa/groups of

the ‘control’ site. The corresponding proportions pertaining to the periods between the two fires and the post-‘second fire’ up to 26 November 1990 were also similar, although the pre-‘first fire’ proportions were significantly higher than those of the post-‘second fire’ (Z = 2.085; n = 8, n = 20; P < 0.05). The three pre- and post-fire data sets for the Shannon-Wiener, Margalef and Pielou community indices (expressed for families) were not significantly different. Thus, the diversity and richness of coleopteran families, and the extent of evenness in the distribution of individuals among them, had not changed following the fires. This conclusion is supported by the observation that 11 of the 31 families trapped (35.5%) were present in all pre- and post-fire samples, while another 13 families (41.9%) were detected after each fire or after the second. Of these 24 families, 83.3% were also observed at the ‘control’ site. Among the remaining seven ‘minor’ families (22.6% of total), a solitary specimen of family Zopheridae was trapped exclusively at the ‘burnt’ site, whereas the families Ptiliidae, Throscidae, Coccinellidae, Mycetophagidae, Colydiidae and Aderidae were each represented by only one to two specimens at the ‘control’ site (Tables 2 and 3).

Table 4 Statistics on activity of total Coleoptera, of ‘major’ families site between 26 March 1985 and 26 February 1990

and Management

3.6 + 0.9

9.6 + 2.8 ab

7.3 + 2.3 a

11.4 + 2.4 a 11.2 f 1.6 a 4.2 + 1.7 b

7.4 + 1.8 a 9.2 + 1.3 a

a Each composite sample consisted of the pool of 20 weekly trap collections and each annual set of samples four seasons. ’ Standard error. ’ The means in each column designated by different letters differ significantly at P < 0.05.

included

1.2 + 0.4 b

observations

from all

23.00 22.81 20.44 21.75 19.50 23.06 20.68

a n denotes no. of observations. b Levels of significant differences

Total Coleoptera Staphylmidae Nitidulidae Leiodidae Minor families Predators Decomposexs/ fungus feeders

are ‘P < 0.05,

16.52 16.57 17.28 16.89 17.55 16.50 17.20

1.572 1.513 0.766 1.185 0.475 1.591 0.845

NS NS NS NS NS NS NS

~.l-.--

26.11 25.82 26.42 21.39 25.18 25.89 26.00

* * P < 0.01, or NS. not significant.

-

21.15 21.50 20.72 27.52 22.40 21.45 21.30

-

1.227 1.077 1.410 1.530 0.694 1.098 1.162

NS NS NS NS NS NS NS

11.85 11.97 12.90 14.07 13.92 12.00 12.62

21.12 20.81 18.50 15.56 15.93 20.75 19.18

*

’ * * NS NS NS

- 1.908 NS

-2.543

- 2.696 -2.571 - 1.628 - 0.435 - 0.587

Table 5 Significance of differences between pre- and post-fire data sets for activity (expressed as differences between contemporaneous fire treatment and control observations) over 7 day period between 26 March 1985 and 26 November 1990 of seven categories of Coleoptera at the litter surface of the study site burnt on 15 October 1985 and 3 November 1988 -Category of Mean rank Z-statistic Mean rank Z-statistic Mean rank Z-statistic Coleoptera and and Pre-first fire Between fires Between fires Post-second fire Pre-first fire Post-second fire significance h significance (n=8)” (n = 27) (n=27) (n=20) (n=8) (n = 20)

3

Pd

2

% 8 F B !i

d

2

$ 1 E T

2

N.G. Collett,

F.G. Neumann/Forest

Ecology

Coleoptera (Figs. 3(a)-3(g)), excepting the fungusfeeding ‘minor’ family Lathridiidae. Immediately after the ‘second fire’, from 3 November to 20 Decem-

and Management

76 (1995)

69-85

81

ber 1988, 200 specimens of Aridius sp. (representing 97.6% of the total lathridiid collection during the entire 5.7 year study period) were trapped at the Cd)

(a) 660 1

Total -

Fire

Fire

1

2 2300

Fire +

Fire i

Fire

Fire

Leiodidae --oBsmu --*--conuo,

Coleoptera Bum (e)

(b) 466

Fire

"1

Minor +

families 9”m

Stapbylinidae Bum,

Fire

---+---

Conm,

t

-

Nitidulidae Bum,

---t-e-

conuol

k?) Fi,re

“1

Deeomposers Bwri

P

6 A WSSu 1995

AWSSuAWSSuA 1996

1987

WSSu 1988

AWSSuAWS 1989

1990

1985

1966

1987

1966

1969

1996

Fig. 3. Temporal variations in monthly activity of total Coleoptera (a), ‘major’ families (b-d), the pool of 28 ‘minor’ families (e), total predators (f) and total decomposers (9, from 26 March 1985 to 26 November 1990 at the litter surface of the site burnt on 15 October 1985 and 3 November 1988 compared with variations at the ‘control’ site (A, autumn; W, winter; S, spring; Su, summer).

82

N.G. Collett,

F.G. Neumann/Forest

‘burnt’ site compared with only one specimen of that species at the ‘control’ site (Table 2). The two-dimensional chi-square tests on the preand post-fire species totals (Table 3) indicated that neither fire had affected the proportions of species totals trapped at the ‘burnt’ site relative to the ‘control’ site. The three-dimensional chi-square tests for mutual independence among the two study sites, the two fire treatments and the species frequency for each of the ‘major’ families and the pool of ‘minor’ families were also insignificant, implying that: (1) no interactions had occurred among these three variables; (2) there were no cumulative effects as a result of the ‘second fire’ on the species profiles of the Staphylinidae, Nitidulidae, Leiodidae and the pool of the ‘minor’ families.

4. Discussion There is presently no comparable study available in the literature on the likely impact of two short-rotation prescribed low-intensity fires in spring on surface-active families and species of Coleoptera in Australian native forest. Previous Australian studies on prescribed fire in relation to the Coleoptera have been cursory, as they have (1) generally considered only the consequences of single fires, and have lacked details on fire behaviour and pre- and post-fire fuel loads (e.g. Springett, 1979; Campbell and Tanton, 1981; Abbott, 1984; Abbott et al., I984), (2) been based on nil or insufficient pre-fire samples (e.g. Majer, 1984), (3) treated the Coleoptera only at ordinal level (e.g. Neumann and Tolhurst, 1991; Collett et al., 1993) (4) examined only the superfamilies Caraboidea and Histeroidea, and the families Curculionidae and Staphylinidae in terms of absence or presence after fire (e.g. Springett, 1976), and lacked statistical tests of significance (e.g. McNamara, 19.55). The study showed that the litter layer of dry sclerophyll eucalypt forest of the type studied is inhabited by a wide range of surface-active Coleoptera, including the 31 families and 105 species identified in the present study. In the absence of fire (as at the ‘control’ site) the activity of the adults of this fauna was shown to remain fairly constant over consecutive years, suggesting that a high degree of

Ecology

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76 (199516945

stability exists among litter-frequenting Colepptera. This conclusion is consistent with that of studies in the same unburnt forest by Neumann and Tolhurst (1991) and Collett et al. (1993) which detected little variation in adult activity among 29 ordinal and sub-ordinal arthropod taxa. As pitfall trapping was used in all of these studies, it appears that this sampling technique gives consistent results in undisturbed forest with respect to temporal activity at the litter surface of arthropods generally and of Coleoptera in particular. Pitfall trapping therefore seems suitable for detecting shortand long-term effects of major ecological disturbances, such as prescribed fire, on surface-active adult Coleoptera. Any limitations in sampling due to the possible inappropriateness of pitfall trapping for some taxa (e.g. the Bostrichidae, Colydiidae and Tenebrionidae which are largely confined to decaying wood systems), or as a result of the relatively small receptive surface area of the traps, would apply equally to the fire-treated and ‘control’ study sites. Moreover, pitfall trapping is unlikely to compromise the analysis of the present study as this concerned sampling over time for fire effects on epigeal Coleoptera and did not aim at a complete census of the coleopteran fauna. The non-significant variations in annual activity of the five broad taxonomic groups at the ‘control’ site over the 5.7 year study period (Table 4(a)) suggests that the epigeal coleopteran populations had been free of large local random protracted effects that could have caused substantialiy different longterm trends in abundance at the ‘control’ site or by inference at the nearby ‘burnt’ site. It therefore appears that the ‘independence assumption’ of the BACI design was satisfied (Stewart-Oaten et al., 1986). As the slopes of the regression lines for ‘control/impact’ differences over time were near zero for all parameters (except for staphylinid activity at the ‘burnt’ site), it also appears that the ‘constancy assumption’ of the BACI design was satisfied, thereby strengthening the validity of the results of the Mann-Whitney U-tests performed on the pre- and post-fire data sets. As a strong seasonal trend was identified among the ‘major’ and ‘minor’ families, sampling for fire effects on taxon richness should ideally be done contemporaneously (as also stipulated by Stewart-

N.G. Coilett,

F.G. Neumann/Forest

Oaten et al., 1986) and over similar time frames and seasons for valid pre- and post-fire comparisons. The pre-first fire statistics for family and species totals (Table 3) correspond to only 7 months of sampling, compared with a 3 year ‘in between fires’ period and a 2 year post-‘second fire’ period. Thus, the much higher family/species totals recorded for the two post-fire periods (Table 3) do not necessarily imply that a substantial increase in family/ species richness had occurred as a result of the fires. The results of the chi-square tests performed on the pre- and post-fire collections of the Coleoptera from the ‘control’ and ‘burnt’ sites provide better indications of the fire effects on family/species richness. As the null hypothesis of a ‘nil effect’ was accepted in all of these tests, it appears that neither the fires on their own, nor their cumulative effect, had significantly altered the family/ species profiles of the Coleoptera. This conclusion is supported by the insignificant results of the Mann-Whitney U-tests on pre- and post-fire data sets based on families for the Shannon-Wiener, Margalef and Pielou community indices. While the family/species richness remained unchanged, the ‘first fire’ in October 1985 appears to have caused short-term upsurges in activity within 2 years of that fire among the ‘major’ families Nitidulidae and Leiodidae and among the pool of the 28 ‘minor’ families and of total decomposers (Figs. 3&3(e) and 3(g)). In contrast, the ‘second fire’ in November 1988 induced a significant reduction in the proportions of Coleoptera among total surfaceactive arthropods, and in activity among total Coleoptera and predators (Table 5). These drops in activity, measured relative to the pre-fire sampling period in 1985, would appear to be linked to the concomitant significant decline in activity observed among the Staphylinidae, which was the largest of all the families trapped (Table 2) and therefore also constituted most of the predators in trap collections. The observed reduction in staphylinid activity during the post-‘second fire’ period relative to the pre-fire period appears to have occurred evenly over the species range as indicated by the non-significant chi-square test on the species profile and by the observation that on the ‘burnt’ site all of the six staphylinid species detected before the ‘first fire’ were trapped after the ‘second fire’ (Table 3). While

Ecology

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76 (1995)

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83

predator activity had decreased following the ‘second fire’, the activity of the decomposers/fungus feeders remained unchanged (Table 5). This indicated that the coleopteran component of the arthropod complex contributing to the decomposer cycle in the litter of the forest type studied is unlikely to be affected by two short-rotation low-intensity fires in spring. The short-term upsurge of the undescribed lathridiid Aridius sp. immediately after the ‘second fire’ might be due to rapid growth of bacteria and/or fungi, especially slime moulds (Myxomycetes) and moulds (Phycomycetes and Ascomycetes) which are a known food source for many insects including lathridiids. A Swedish study has reported a similar upsurge in lathridiid activity immediately after fire in response to an increase in slime moulds, and moulds (Wikars, 1992). Aridius sp. therefore appears to be opportunistic, taking advantage of a set of favourable conditions to increase its activity, rather than forming a stable part of the coleopteran litter fauna. Notwithstanding short-term increases in activity by a few coleopteran taxa in response to two shortrotation low-intensity prescribed fires in spring, the study indicates that any such fire regime may cause a decline in activity of the ‘major’ family Staphylinidae, and with it a decline in activity among total Coleoptera and predators at litter level. Burning in autumn instead of spring is not expected to alleviate this impact as staphylinid activity was found to be similar during autumn and spring (Table 4(b)). However, as species richness was not affected, it is likely that the staphylinid populations in areas burnt in spring will return to pre-fire levels in due course without unacceptable adverse impacts occurring, provided no further fires are applied or occur naturally in the short term.

Acknowledgements We thank Research Officer K. Tolhurst for providing data on fire behaviour and fuel load statistics, and Research Manager Dr. D.W. Flinn for his encouragement and support. It is also a pleasure to acknowledge T. Weir and his staff at the Australian National Insect Collection, Division of Entomology,

84

N.G. Collett,

F.G. Neumann/Forest

CSIRO Canberra, for providing authoritative identifications of many of the beetle species trapped.

References Abbott, I., 1984. Changes in the abundance and activity of certain soil and litter fauna in jarrah forest of Western Australia after a moderate intensity fire. Aust. J. Soil Res., 22: 463-469. Abbott, I., van Heurck, P. and Wong, L., 1984. Response to long-term fire exclusion: physical, chemical and fauna1 features of litter and soil in a Western Australian forest. Aust. For., 47: 237-242. Booth, R.G., Cox, M.L. and Madge, R.B., 1990. BE Guides to Insects of Importance to Man. 3. Coleoptera. International Institute of Entomology, The University Press, Cambridge. Byram, G.M., 1959. Combustion of forest fuels. In: K.P. Davis (Editor), Forest Fire, Control and Use. McGraw-Hill, New York. Campbell, A.J. and Tanton, M.T., 1981. Effects of fire on the invertebrate fauna of soil and litter of a eucalypt forest. In: A.M. Gill (Editor), Fire and the Australian Biota. Australian Academy of Science, Canberra, pp. 216-241. Christensen, P. and Abbott, I., 1989. Impact of fire in the eucalypt forest ecosystem of southern Western Australia: a critical review. Aust. For., 52: 103-121. Collett, N.G., Neumann, F.G. and Tolhurst, K.G., 1993. Effects of two short-rotation prescribed fires in spring on surface active arthropods and earthworms in dry sclerophyll eucalypt forest of west-central Victoria. Aust. For., 56: 49-60. Department of Conservation and Environment (Victoria), 19891991. Annual Reports, 1989-90, 1990-91. Department of Conservation, Forests and Lands (Victoria), 19831989. Annual Reports, 1983-84,1984-85,1985-86,1986-87, 1987-88, 1988-89. Department of Conservation, Forests and Lands (Victoria), 1986. Policy on Fuel Reduction Burning. 8 pp. Forbes, S.J. and Ross, J.H., 1988. A Census of the Vascular Plants of Victoria, 2nd edn. Department of Conservation, Forests and Lands, Melbourne. Forests Commission Victoria, 1983. Annual Report 1982-83. Green, R.H., 1993. Application of repeated measure designs in environmental impact and monitoring studies. Aust. J. Ecol., 18: 81-98. Greenslade, P.J.M., 1964. Pitfall trapping as a method for studying populations of Carabidae. J. Anim. Ecol., 33: 301-310. Greenslade, P. and Greenslade, P.J.M., 1971. The use of baits and preservatives in pitfall traps. J. Aust. Entomol Sot., 10: 2.53-260. Greig, P.J., 1986. Forest policy developments in Victoria. Aust. For., 49: 197-202. Hurlbert, S.H., 1971. The non-concept of species diversity. A critique and alternative parameters. Ecology, 52: 577-586.

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Johnston, J.F., McKittrick, D.J., Flier, D.W. and Brown, H.G., 1983. Fire protection and fuel reduction burning in Victoria. Report to the Minister of Forests, Forests Commission, Victoria, April 1983. Lawrence, J.F. and Britton, E.B.. 1991. Coleoptera. In: ID Naumann @ditor), The Insects of Australia. Division of Entomology, CSIRO, Melbourne Ilniversity Press, Carlton, pp. 543-683. Luke, R.H. and McArthur, A.G., 1978. Bushfires in Australia, A.G.P.S., Canberra, 359 pp. Majer, J.D., 1984. Short-term response of soil and litter invertebrates to a cool autumn bum in Janah (Euca&ar marginural forest in Western Australia. Pedobiologia, 26: 229-247. McNamara, P.J., 1955. A preliminary investigation of the fauna of humus layers in the jarrah forest of Western Australia. Commission of Australia, Forest and Timber Bureau, Leaflet No. 71, 16 pp. Mount, A.B., 1972. The Derivation and Testing of a Soil Dryness Index Using Run-off Data. Bulletin No. 4, Tasmania Forestry Commission. Neumann, F.G., 1987. Introduced bark beetles on exotie trees in Australia with special reference to infestations of fps grandicollti in pine plantations. Aust. For., 50: 166178. Neumann, F.G. and Tolhurst, K., 1991. Effects of fuel reduction burning on epigeal arthropods and earthworms in dry sclerophyll eucalypt forest of west-central Victoria. Aust. J. Ecol.. 16: 315-330. New, T.R., 1987. Insect conservation in Australia: towards national ecological priorities. In: J.D. Majer (Editor), The Roie of Invertebrates in Conservation and Biological Survey. Proceedings of the 18th Scientific Conference of the Australian Entomological Society, Perth, Australia, pp. 5-20, Pielou, E.C., 1966. The measurement of diversity. in different types of biological collections. J. Theor. Biol., 13: 131-144. Poole, R.W., 1974. An Introduction to Quantitative Ecology. McGraw-Hill, New York. Pratten, C., 1985. Policies of the NSW voluntary conservation movement regarding fuel reduction burning. Paper to Symposium on Fighting Fire with Fire. Graduate fchool of Environmental Science, Monash University, Clayton, Vie. Rawson, R.P., Billing, P.R. and Duncan, S.F., 1983. The 1982-83 forest fires in Victoria. Aust. For., 46: 163-172. Southwood, T.R.E., 1978. Ecological Methods, with Particular Reference to the Study of Insect Populations. Methuen, London. Springett, J.A., 1976. The effect of prescribed burning on the soil fauna and on litter decomposition in Western Australia forests. Aust. J. Ecol., 1: 72-82. Springett, J.A., 1979. The effects of a single hot summer fire on soil fauna and on litter decomposition in jarrah-(Eucalyptus marginatu) forest in Western Australia. Amt. J. EcoI., 4: 279-291. Stewart-Oaten, A., Murdoch, W.W. and Parker, K.R., 1986. Environmental impact assessment: ‘pseudorepliiation’ in time’? Ecology, 67: 929-940. Tolhurst, K.G. and FIinn, D.W., 1992. Ecological impacts of fuel reduction burning in dry sclerophyll forest: first progress

N.G. Collett, F.G. Neumann/ Forest Ecology and Management 76 (1995) 69-85 report. Department of Conservation and Environment (Victoria), Research Report 349, 270 pp. Tolhurst, KG. and Turvey, N.D., 1992. Effects of bracken (Pteridium esculenrum) (Forst.f.1 (Cockayne) on eucalypt regeneration in west-central Victoria. For. Bcol. Manage., 54: 45-67.

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Wikars, L.O., 1992. SkogsbrZnder och insekter. Entomol. Tidskr., 113: l-11. Zar, J.H., 1984. Biostatistical Analysis. Prentice Hall, London.