Effects of two silvicultural practices on soil fauna
abundance in a northern hardwood forest,
Québec, Canada
Jean-David Moore1, Rock Ouimet1, Claude Camiré2, and Daniel Houle1
1Direction
de la recherche forestière, Forêt Québec, ministère des Ressources naturelles du Québec, 2700 rue
Einstein, Sainte-Foy, Québec, Canada G1P 3W8 (e-mail: [email protected]); 2Centre de
recherche en biologie forestière, Faculté de foresterie et de géomatique, Université Laval, Sainte-Foy, Québec,
Canada G1K 7P4. Received 30 March 2001, accepted 5 October 2001.
Moore, J.-D., Ouimet, R., Camiré, C. and Houle, D. 2002. Effects of two silvicultural practices on soil fauna abundance in a
northern hardwood forest, Québec, Canada. Can. J. Soil Sci. 82: 105–113. Soil fauna play a key role in soil fertility and productivity of forest ecosystems and represent an important base of terrestrial food chains. The impact of forest management on soil
fauna should be considered when sustainable forest management and conservation of biodiversity are desired. We evaluated the
impact of selective cutting and strip clearcutting on soil fauna abundance in a northern hardwood forest of the Lower Laurentians
of Québec. Twelve years after strip clearcutting, the abundance of carabid beetles (Coleoptera: Carabidae), collembolans
(Hexapoda: Collembola) and snails (Stylommatophora: Sigmurethra) was greater in the strip clearcuts than the adjacent undisturbed strips. Snails and millipedes (Polydesmida: Polydesmidae) were more abundant in the selective cuts 6 to 8 yr after treatment. Spiders (Arachnida) were the only organism whose abundance was lower in the selective cuts than in the adjacent
undisturbed forest. No significant negative effect of the silvicultural treatments was noted for the abundance of other caught organisms. This one-season sampling suggests there are few negative impacts associated with low intensity selective cutting and strip
clearcutting on the abundance of soil fauna in this northern hardwood forest stands 6 to 12 yr after harvest.
Key words: Salamander, arthropod, shrew, northern hardwood, selective cutting, strip clearcutting
Moore, J.-D., Ouimet, R., Camiré, C. et Houle, D. 2002. Effets de deux pratiques sylvicoles sur l’abondance de la faune du sol
dans une érablière, Québec, Canada. Can. J. Soil Sci. 82: 105–113. La faune du sol joue un rôle primordial en ce qui concerne
la fertilité du sol et la productivité des écosystèmes forestiers et constitue un maillon important de la chaîne alimentaire terrestre.
Les impacts des activités forestières sur les organismes du sol devraient donc être considérés dans un contexte d’aménagement
forestier durable et de conservation de la biodiversité. Nous avons évalué l’impact de coupes de jardinage et de coupes totales par
bandes sur l’abondance de la faune du sol dans une érablière des basses Laurentides de la région de Québec. Douze ans après la
coupe totale par bandes, l’abondance des carabidés (Coleoptera: Carabidae), des collemboles (Hexapoda: Collembola) et des
escargots (Stylommatophora: Sigmurethra) était plus élevée dans les bandes coupées comparativement aux bandes adjacentes
non-perturbées. Les escargots et les millipèdes (Polydesmida: Polydesmidae) étaient plus abondants dans les forêts jardinées 6 à
8 ans après traitement comparativement à celles témoins. L’araignée (Arachnida) représentait le seul organisme dont l’abondance
était plus faible dans les forêts jardinées comparées aux secteurs témoins. Aucun effet significatif des traitements sylvicoles n’a
été décelé sur l’abondance des autres organismes capturés. Cette campagne d’échantillonnage d’une saison suggère que les coupes
de jardinage de faible intensité et les coupes totales de faible superficie dans ce type de forêt engendre peu d’impacts négatifs sur
l’abondance de la faune du sol 6 à 12 ans après les traitements.
Mots clés: Salamandres, arthropodes, musaraignes, érablière, coupe de jardinage, coupe par bandes
Arthropods play a key role in soil fertility and the
productivity of forest ecosystems by taking part in
nutrient turnover (Reichle et al. 1969; Crossley 1977;
Edmonds and Eglistis 1989). Moreover, arthropods and
other soil fauna represent an important base of terrestrial
food chains and are vital to the survival of many forest
wildlife species (South 1980; Vaughan and Weil 1980;
Kremen et al. 1993). Despite their importance, few studies
have investigated the impact of forest management on soil
arthropods in northeastern United States and Canada
(Jennings et al. 1984; Bird and Chatarpaul 1986; Duchesne
et al. 1999) in comparison to salamanders (Caudata) (c.f.
deMaynadier et Hunter 1995, 1998; Messere and Ducey
1998; Satter and Reichenbach 1998; Harpole and Haas
1999) and shrews (Soricidae) (c.f. Kirkland 1990; Mitchell
et al. 1997; Menzel et al. 1999). Understanding the impacts
of forest management on soil fauna is essential to achieve
sustainable forest management and conservation of biodiversity. We evaluated the impact of selective cutting and
strip clearcutting on the relative abundance of soil arthropods, salamanders, shrews, and terrestrial molluscs in a
sugar maple forest of the Lower Laurentians in Québec. We
tested the hypothesis that these forest management practices
would have a minimal impact on the abundance of soil
fauna.
To our knowledge, this study is the first to report the
effects of forest practices on the soil fauna in northern hardwood forests of Canada.
105
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CANADIAN JOURNAL OF SOIL SCIENCE
MATERIALS AND METHODS
Site Description
The Duchesnay forest station (46°57′N, 71°40′W) is located
approximately 50 km northwest of Québec City, Québec,
Canada. The elevation varies between 270 and 390 m, and
the average slope is approximately 10%. The mean annual
temperature is 3.4°C and annual precipitation is 1300 mm.
According to the Canadian system of soil classification
(Canada Soil Survey Committee 1992), the soil is mainly an
Orthic Ferro-Humic Podzol and pertains to the stony loamy
sand of the Sainte-Agathe series (Raymond et al. 1976). The
humus is of mor type, and the main surface deposit is a very
acidic and stony till on an underlying bedrock made of granite and syenite.
Two silvicultural practices were used in this study: selective cutting and strip clearcutting. In the selective cutting
and strip clearcutting areas, the forest was dominated by
mature (85 to 130 yr old) sugar maples (Acer saccharum
Marsh.) in association with yellow birch (Betula
alleghaniensis Britt.) and American beech (Fagus grandifolia Ehrh.) forming a closed forest canopy. Strip clearcutting
was implemented in 1983 and 1984. The strips were 60 m
wide and approximately 1000 m long, one cut strip alternating with an uncut strip, repeated two times. Regeneration
sprouted rapidly in strip clearcuts making up a dense stand
of pin cherry (Prunus pensylvanica L.f.) and yellow birch of
3.5 m height in 1996.
In the selective-cut plots, small gaps were created in 1988
and 1990. Characteristics of the selective cut sites may be
found in Majcen and Richard (1995) and Majcen (1996). In
brief, each one of the two selective-cut plots included a
block of 2 ha in the treated area and a block of 1 ha in the
untreated area (control). Whereas the strip clearcutting areas
were adjacent to each other, the distance between the two
selective cutting areas was approximately 9 km. Basal area
(BA) in the control plots in 1988 and 1990 was 26.4 and
24.7 m2 ha–1, respectively. The BA before and after cuts was
25.6 and 17.5 m2 ha–1 in 1988, and 24.9 and 18.8 m2 ha–1 in
1990. The intensity of the selective cutting was 32% in 1988
and 25% in 1990. No major forest harvesting occurred in the
study area before the present forest operations for at least
150 yr.
Coarse woody debris (CWD) were more abundant in
treated forests than in control sites given that non-commercial woody biomass was left on the plots after selective and
strip cutting. However, there was no attempt to obtain a
quantitative assessment of CWD left on the sites.
Experimental Set-up
Pitfall arrays were established in June 1996 in two repetitions of the two silvicultural practices, each one associated
with a control forest site, for a total of eight arrays. Six pitfall traps were installed per array, for a total of 48 pitfall
traps. No drift fences were used.
In the strip clearcuts, the six pitfall traps were distributed
every 15 m along a transect located at the centre (30 m) of
the strips. In selective cutting areas, traps were placed systematically at the four corners of the plots and in the middle
of two opposite sides, with a minimal distance of 50 m
between each trap. The pitfall traps were the “Multi-Pher”
type (Jobin and Coulombe 1988). This slightly conical trap
has a 20.5-cm height with a diameter of 10.0 cm at the base
and 13.0 cm at the top. A funnel of 4.0 cm diameter at the
base were located in the top part of the trap. Some moist
leaves were placed in the pitfall traps. No baits or preservatives were used. Soil fauna were collected every 2 d, from
26 June to 5 July, from 12 July to 16 August, and from 23
August to 2 September 1996, for a total sampling time of 57
d. Salamanders were identified in situ and released at nearby sites that were >2 km from our study areas. Shrews and
invertebrates were transported separately to the laboratory
and frozen for later identification. Collected salamanders
and shrews were identified to species, while invertebrates
were identified to order or family.
Statistical Analyses
To check for dependence between trap captures, a variographic analysis was conducted on the total captures for
each organism group using the GEOEAS software (Glund
1991). To verify independence between traps, the strip
clearcutting experiment was used for spatial analyses
because the distances between two adjacent traps were
smaller (15 m) than those in the selective cutting experiment
(≥ 50 m). Organism abundance was pooled weekly, and we
analyzed the effect of the two silvicultural practices separately by the regression of generalized linear models. The
GENMOD SAS procedure was used by specifying the
Poisson distribution, the standardization (option DSCALE)
and the logarithmic function as model options (SAS
Institute, Inc. 1989). This procedure was used to test for
treatment (cut vs. control) and time (sampling week) effects
on the abundance of each organism group.
RESULTS AND DISCUSSION
Spatial Analyses
The results of the spatial analyses show independence of
traps for most organisms. The variance associated with
organism abundance (total captures) was constant, or
increased with distance between traps, suggesting that number of captures at one location did not influence the number
of captures elsewhere in the strips (Fig. 1). Each trap was
therefore considered as an independent sample for a given
treatment. However, three exceptions occurred. The variability of carabid beetle captures in the clearcut strips
showed a periodic cycling pattern within a distance of 30 m.
The semivariogram of carabid beetle abundance suggests
that captures at traps located at a 15-m distance influenced
each other. Captures of crickets in the clearcut strips showed
a decrease in variance with distance between traps. This pattern indicates that as sampling points are closer to each
other, variability increases between samples, possibly
because of the captures themselves. A similar pattern was
observed for salamanders in the control strips. These results
suggest that 15 m between traps was too close for carab beetles, crickets and salamanders. Results from the strip
clearcutting analyses concerning these three groups of
MOORE ET AL. — SOIL FAUNA AND FOREST CUTTING
107
Fig. 1. Semivariograms of total number of captures of spiders, collembolans, carabid beetles, millipedes, snails, slugs, shrews, crickets, and
salamanders in the clearcut and control strip areas. Semivariograms stratify variances by the distance (lag) separating each pair of sampling
points.
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CANADIAN JOURNAL OF SOIL SCIENCE
Table 1. Mean capture number (± standard error) by trap by week and probability value of the effect of treatment and sampling period on soil organisms collected during summer 1996 in selective and strip cutting harvest techniques
P value
Capture number
(no. by trap–1 wk–1)
Forest harvest
Organism
Strip clearcutting
Selective cutting
Treatment ×
Treated
Control
Treatment
Sampling period
sampling period
Spider
Millipede
Carabid beetle
Collembolan
Cricket
Slug
Snail
Salamander
Shrew
1.77 (0.17)
2.03 (0.21)
4.90 (0.52)
51.28 (4.56)
0.79 (0.13)
0.94 (0.21)
0.37 (0.07)
0.08 (0.03)
0.18 (0.05)
1.59 (0.16)
2.40 (0.27)
2.86 (0.31)
32.15 (2.65)
0.64 (0.13)
1.01 (0.19)
0.26 (0.06)
0.10 (0.04)
0.26 (0.06)
0.359
0.496
<0.001
<0.001
0.140
0.593
0.028
0.388
0.174
<0.001 (Q: 0.042)
0.222
<0.001 (L: <0.001)
<0.001 (Q: <0.001)
0.079
<0.001 (Q : <0.001)
0.233
0.159
<0.001 (C : <0.001)
0.614
0.396
0.893
0.663
0.122
NE
0.206
NE
NE
Spider
Millipede
Carabid beetle
Collembolan
Cricket
Slug
Snail
Salamander
Shrew
1.21 (0.12)
2.14 (0.27)
3.51 (0.35)
35.65 (4.24)
1.58 (0.18)
0.53 (0.09)
0.17 (0.05)
0.14 (0.04)
0.32 (0.07)
1.77 (0.21)
1.27 (0.17)
3.69 (0.50)
39.83 (3.83)
1.81 (0.22)
0.46 (0.09)
0.08 (0.03)
0.21 (0.05)
0.33 (0.08)
0.011
0.002
0.756
0.254
0.306
0.074
0.011
0.124
0.223
0.188
<0.001 (Q: <0.001)
<0.001
<0.001 (Q: <0.001)
0.047 (L: <0.001)
<0.001 (L: <0.001)
0.195
0.014 (Q: 0.023)
0.344
0.229
0.445
0.033
0.823
0.847
0.199
NE
NE
0.459
Note: NE = non estimable, L = linear, Q = quadratic and C = cubic effects.
organisms should therefore be considered with caution,
given the dependency pattern between capture locations.
Given the large distance between traps in the selective cutting experiment (≥ 50 m), capture samples should be independent.
The semivariograms in Fig. 1 also show that variance
associated with spider, collembolan, snail, carabid beetle
and shrew abundances was lower in the control strips compared to the clearcut strips. This may reflect changes in
organisms at the species level, in behaviour, food source or
habitat following forest cutting.
Salamanders
Four species of salamander were collected, totalling 52 captures: Plethodon cinereus (n = 41); Ambystoma maculatum
(n = 5); Eurycea bislineata (n = 5); and Notophthalmus viridescens (eft stage) (n = 1). Nearly 80% of captures were redbacked salamanders (Plethodon cinereus), which was
expected given its wide distribution (Grobman 1944; Smith
1963; Bider and Matte 1994) and relative abundance in
forests of eastern North America (Burton and Likens
1975a). The low capture rate of salamanders in our study
(0.02 salamander d–1 trap–1), compared with Bennett et al.
(1980) and Mitchell et al. (1997) (0.07 and 0.05, respectively), cannot be attributed only to the trapping method.
Indeed, McLeod and Gates (1998) also noted a low capture
rate (0.03), in spite of the use of drift fences twinned with
pitfall traps, which is more effective than the use of pitfall
traps alone (Adams and Freedman 1999). A drought period
could not explain the low capture rate of salamanders in our
study since precipitation (and consequently soil humidity)
was not a restricting factor during the sampling period (July
1996 = 230 mm whereas the average from 1989 to 1997 =
142 mm; August 1996 = 120 mm whereas the average from
1989 to 1997 = 127 mm). The most probable explanation for
the low capture rate is reduced abundance, most likely related to the strong acidity of the forest floor at Duchesnay
(pHLF = 3.23 and pHH = 3.05) (Houle et al. 1997). Degraaf
and Rudis (1990) also observed a low capture rate of salamanders (0.03) on acidic soil (pH varying from 3.4 to 3.9).
This is in agreement with studies showing that salamanders
prefer neutral or basic environments over acidic areas
(Vernberg 1955; Mushinsky and Brodie 1975; Wyman and
Hawksley-Lescault 1987; Wyman and Jancola 1992).
Our data suggest there was no effect on salamander abundance 12 yr after strip clearcutting (Table 1). Harper and
Guynn (1999) also reported a quick recovery of the salamander population after clearcutting. Other studies, however, have noted a decline in salamander population following
clearcutting for several decades (Ash 1997; Mitchell et al.
1997; Pough et al. 1987; Petranka et al. 1994).
Selective cutting had no effect on salamander abundance
at Duchesnay 6 or 8 yr after treatment (Table 1). Messere
and Ducey (1998) reported that low intensity selective cutting (gaps from 22 to 94 m2) did not affect red-backed salamander abundance 1 yr after cutting. Harpole and Haas
(1999), however, noted that salamanders were less abundant
in selective cuts compared with control forests 1 to 3 yr after
cutting. Residual BA in their study (selective cutting: 4–7
and 12–15 m2, scattered seed tree cutting: 3–4 m2) were
much lower than the residual BA at Duchesnay after 6 and
8 yr (~ 21 m2). In an other study, Ross et al. (2000) noted a
greater abundance of salamanders with an increase in residual BA; a BA > 15 m2 ha–1 seems to be more favourable to
salamanders. It appears that conservation of a high canopy
closure with selective cutting or the proximity of uncut forest to strip clearcutting may influence salamander abundance in those stands soon after harvest.
MOORE ET AL. — SOIL FAUNA AND FOREST CUTTING
The abundance of CWD observed in the treated areas, as
well as the high forest canopy closure, could also have contributed to a recovery or maintenance of salamander populations after cutting by offering shelter for them. Indeed,
Brooks (1999) noted that number of red-backed salamanders was correlated with the amount of CWD and vegetation
density ≥ 1 m. Also, Butt and McComb (2000) observed an
increase in salamander abundance with an increase in CWD.
Shrews
Three species of shrews were collected, totalling 83 captures:
Sorex cinereus (n = 66); Blarina brevicauda (n = 10); and
Sorex hoyi (n = 7). The masked shrew (Sorex cinereus) is one
of the most abundant small mammals in the forests of North
America (Kirkland 1977; Mitchell et al. 1997; McLeod and
Gates 1998; Menzel et al. 1999). The pygmy shrew (Sorex
hoyi) is likely to be designated as threatened or vulnerable in
the province of Québec (Collin et al. 1996). Our capture rate
of shrews (0.03 shrew d–1 trap–1) is similar to that of Menzel
et al. (1999), who used pitfall traps alone (0.03 shrew d–1
trap–1), and to Mitchell et al. (1997) and McLeod and Gates
(1998) who used drift fences combined with pitfall traps
(0.05 and 0.03 shrew d–1 trap–1, respectively).
Our results showed no difference in the abundance of
shrews between treated and control forests 6 or 12 yr after
selective or strip cutting, respectively (Table 1). Other studies also showed that shrew populations remained stable after
forest cutting (Kirkland 1977; Clough 1987; Brooks and
Healy 1988; Mitchell et al. 1997).
Invertebrates
Four classes of invertebrates were collected during the
study: Arachnida (including Phalangida and Araneida);
Diplopoda (including Polydesmida); Gastropoda (including
Stylommatophora, Sigmurethra); and Hexapoda (including
Coleoptera, Collembola, Orthoptera) (Table 1).
Class Arachnida
The abundance of spiders (Arachnida) at Duchesnay was
lower in selective cuts compared to control areas (Table 1).
This supports the observations of Duffey (1978), who found
that small modifications in habitat structure can have a significant impact on species composition and relative abundance of spiders. Our result are also in agreement with
DeBakker et al. (2000), who suggested that spider community composition is influenced by density of tree coverage.
We found no difference, however, in spider abundance
between the strip clearcuts and the adjacent undisturbed
strips. This also agrees with Harper and Guynn (1999) who
reported no difference in spider abundance or biomass
among forest stand type or age in the southern
Appalachians. A change in species composition after
clearcutting (McIver et al. 1992) and a different response of
the spider community to the two silvicultural practices could
explain our findings at Duchesnay.
Class Diplopoda
Millipedes (Polydesmida: Polydesmidae) were more abundant in the selective cut than in control areas 6 to 8 yr after
109
cutting (Table 1). In the strip clearcutting area, no difference
in capture rate was noted compared with the control strips.
Marra and Edmonds (1998) observed no difference in
Diplopod abundance 2 to 4 yr after a clearcut. It is difficult
to explain the change in millipede abundance at Duchesnay;
however, the most probable explanation is the retention of a
large portion of the forest canopy (keeping temperatures relatively low and moisture relatively high) combined with the
addition of CWD from selective cutting, which may have
created favourable conditions for those organisms.
Furthermore, field observations indicated that young millipedes were more numerous under CWD than under leaf litter. More sampling and research, however, are necessary to
clearly identify reasons for this change in abundance in
selective cut areas.
Class Gastropoda
Following selective or strip cutting, snails (Stylommatophora,
Sigmurethra) were more abundant in the treated forests than
in the control sites (Table 1). No effect of forest harvesting,
however, was observed on the abundance of slugs
(Stylommatophora, Sigmurethra). In spite of the importance
of snails in the diet and the reproductive success of many
vertebrates (Burton and Likens 1975b; Graveland et al.
1994), few studies have investigated the impact of forest
management on these organisms. Strayer et al. (1986)
observed a quick recovery of Gastropod communities following forest disturbance, attributing this phenomenon to
rapid recovery of the vegetation, and to the small area of disturbed territories, facilitating recolonization from surrounding areas. The abundance of CWD at Duchesnay after
cutting may explain the greater abundance of snails in the
treated forests. Field observations indicated the importance
of CWD for young snail specimens given their abundance
under these micro-sites.
Class Hexapoda
After 12 yr, collembolans (Collembola) were more abundant in the strip clearcuts than in the control strips (Table 1).
Bird and Chatarpaul (1986) also found a greater abundance
of collembolans in a conventional forest harvest (where
CWD are generally more abundant) than in a whole-tree
harvest. In contrast, Huhta (1976), Seastedt and Crossley
(1981), Blair and Crossley (1988), and Addison and Barber
(1997) noted that collembolans abundance remained
unchanged in the first few years after clearcutting. Our finding supports the observations of Addison and Barber (1997),
that collembollans are responding positively to woody
debris left on-site after harvest. This observation suggests
that food availability for collembolans was still high in the
strip clearcuts 12 yr after harvest.
Carabid beetles (Coleoptera: Carabidae) were more
abundant in strip clearcuts than in control strips; however,
there was no difference between selective cut areas and
control areas (Table 1). A general pattern in the change of
carabid beetle abundance after forest cutting could not be
found in the literature. Huhta (1976) noted an increase in
carabid beetle populations early after clearcutting (1 to 3
yr); however, Marra and Edmonds (1998) noted a decrease
110
CANADIAN JOURNAL OF SOIL SCIENCE
Fig. 2. Effect of the sampling week on capture rates of soil fauna. Points were placed at the middle of the sampling week.
MOORE ET AL. — SOIL FAUNA AND FOREST CUTTING
in numbers 2 and 3 yr after clearcutting. Addison et
Barber (1997), Greenberg and Thomas (1995) and
Duchesne et al. (1999) observed no differences in carabid
beetle abundance 2, 7 and 9 yr after forest clearcutting,
respectively. As with collembolans, the greater number of
carabid beetles in strip clearcuts could be explained by an
increase of CWD after cutting, as it generally constitutes a
good shelter (Lindroth 1961–1969; Larochelle 1972). The
presence of additional food (e.g., collembolan) also could
explain the difference in carabid beetle abundance
(Guillemain et al. 1997).
No effect of timber harvest was observed on the abundance of crickets (Orthoptera: Gryllacrididae) (Table 1).
Changes in Capture Rate of Soil Fauna during the Sampling
Period
The capture pattern of the soil fauna during the sampling
period was rather variable (Table 1). However, the temporal
variation of collembolan captures was similar for the treated
and control forest for the two experiments (Table 1; Fig. 2).
Capture rates were low at the beginning and at the end of the
summer, whereas a maximum was noted between 19 July
and 9 August 1996. Blair and Crossley (1988) noted a similar temporal pattern after clearcutting.
Population decline due to removal sampling was not
important in our study (Table 1; Figs. 1 and 2). This is somewhat surprising since the capture rate immediately after
installing pitfall traps is usually higher than subsequent captures (Greenslade 1973).
Only the capture rates of carabid beetles and slugs
decreased in the two silvicultural experiments during the
sampling period. This phenomenon may have been caused
by a reduction in population size attributable to sampling
(see Figs. 1 and 2 for the carabid beetle). To determine the
reason for these changes in capture rates, an array of pitfall
traps at fixed locations should be compared with another
series with variable locations (i.e., with a change of their
location at regular intervals).
The short sampling period (one summer) considered in
this study does not permit strong conclusions about the
effects of forest harvesting on soil fauna. It is possible that a
longer study may yield different results, particularly for the
strip clearcutting experiment, whose perturbations by forest
harvesting were greater than those on selective cut sites.
Indeed, abundance of soil fauna may change during years
following forest harvest due to the reestablishment of initial
conditions (e.g., salamander: Ash 1997).
CONCLUSIONS
This one-season study suggests there are few negative
impacts from low-intensity selective cutting or strip
clearcutting on the abundance of soil fauna in northern hardwood stands 6 to 12 yr after harvest. The few differences in
the abundance of soil organisms between the treated and
control areas may be explained by (1) the presence of uncut
forest near the harvested areas, (2) the retention of the
majority of the forest canopy in the selective cut areas, (3)
the fast recovery of vegetation cover in strip clearcuts, and
(4) the abundance of CWD left after the cuts.
111
ACKNOWLEDGEMENTS
Thanks to B. Drolet, M. Pelletier and D. Johnston for the
invaluable assistance in the identification and preparation of
the soil organisms. Moreover, the collaboration of M. Z.
Majcen and L. Bélanger largely facilitated the realisation of
this study. We also thank J. C. Mitchell and C. A. Harper for
their constructive comments on a former version of the manuscript. This study was part of a research project on forest
soil modelling/biodiversity (project no. 0310 149S) at the
Direction de la recherche forestière, Forêt Québec, ministère
des Ressources naturelles.
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