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7/25/2019 Changes in Soil and Litter Arthropod Abundance Following Three Harvesting and Site Preparation in a Loblolly Pine
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Changes in soil and litter arthropod abundance followingtree harvesting and site preparation in a loblolly pine
(Pinus taeda L.) plantation
Simon B. Birda,*, Robert N. Coulsonb, Richard F. Fisherc
a
Centre for Ecology and Hydrology Bangor, School of Agriculture and Forestry Sciences, University of Wales Bangor,Deiniol Road, Bangor, Gwynedd LL57 5UP, UKbKnowledge Engineering Laboratory, Department of Entomology, Texas A&M University, College Station, TX 77843, USA
cTemple Inland, PO Drawer N, 303 South Temple Drive, Diboll, TX 75941-0814, USA
Received 16 June 2003; received in revised form 5 May 2004; accepted 8 July 2004
Abstract
Soil and litter arthropods are important in many forest ecosystem processes where they help to regulate nutrient dynamics
and soil quality, and are useful bioindicators of ecosystem condition and change. This study was initiated in response to
concerns about possible decline in site productivity due to intensive forestry practices. We investigated the effects of tree
harvesting and site preparation treatments on soil and litter arthropod abundance in a loblolly pine plantation in easternTexas, USA. Using soil and litter cores, we sampled abundance of selected arthropods over two years following tree
harvesting. Response to treatments varied somewhat among arthropod taxa. Acari (mites) and Collembola (springtails), the
numerically dominant taxa in core samples, were initially higher in abundance in less intensive harvesting and site
preparation treatments. However, after 2 years, abundance of these arthropods was comparable in all harvesting and site
preparation treatments. Fertilization with nitrogen and phosphorus had a strong positive effect on abundance of most
arthropod groups in the second year of the study. The recovery of arthropod abundance through time suggests that the
silvicultural practices used did not jeopardize the ecological integrity of the site. The results reported here contrast with
other similar studies which suggests that soil and litter arthropod communities respond differently in different geographic
locations and forest types. Further comparative and extensive studies of this nature are needed therefore for a deeper
understanding of the impacts of forest management practices.
# 2004 Elsevier B.V. All rights reserved.
Keywords: Soil and litter arthropods; Effects of forest management; Soil ecology; Silvicultural practices; Mites; Springtails
1. Introduction
Soil and litter arthropods are important components
of forest ecosystems, and they play a particularly
www.elsevier.com/locate/foreco
Forest Ecology and Management 202 (2004) 195208
* Corresponding author. USDA-ARS Jornada Experimental
Range, New Mexico State University, MSC 3JER, Las Cruces,
NM 88003, USA. Tel.: +1 505 646 4152; fax: +1 505 646.
E-mail address: sb2417@columbia.edu (S.B. Bird).
0378-1127/$ see front matter # 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2004.07.023
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significant role in the process of decomposition. In
addition, these organisms affect porosity, aeration,
infiltration, and the distribution of organic matter
within the soil. Soil and litter arthropods can,therefore, be useful bioindicators of the effects of
land management on nutrient dynamics and site
productivity.
There has been recent concern regarding the
possibility of detrimental ecological effects caused
by forest management practices in the United States.
Silvicultural practices have been placed under
increased scrutiny in respect to their environmental
impacts and effects upon site productivity and
biodiversity (Burger and Zedaker, 1993; Gupta and
Malik, 1996). Tree harvesting and site preparation
practices, as part of intensive management regimes in
North American forests, can lead to significant loss of
nutrients and organic matter from forest ecosystems,
alteration of soil physical properties, significant
disturbance to trophic systems, and overall decrease
in site productivity (Likens et al., 1970; Pritchett and
Wells, 1978,Pritchett and Fisher, 1987; Bormann and
Likens, 1994).
Studies investigating the decline of site productiv-
ity following harvesting and site preparation in
southern US pine (Pinus spp.) plantations have
demonstrated mixed results. In their review, Powerset al. (1990) found that the reports of productivity
decline due to intensive forest management were
inconclusive. However, they suggested that site
preparation treatments causing soil compaction and
organic matter removal pose the greatest risk to site
productivity over successive rotations. The study of
the responses of soil organic matter, trophic system
dynamics, and decomposer communities to forest
management has not been comprehensive across
geographic areas and the wide variety of management
techniques employed in North American forestplantations. This may in part explain the lack of
agreement among results. Field studies designed to
analyze the effects of disturbance and management
practices on soil organic matter, and the biological
processes that regulate soil fertility and nutrient
conservation, are essential for defining sustainable
production systems (Gupta and Malik, 1996).
The goal of this study was to investigate the effects
of silvicultural practices of varying intensity on the
abundance of soil and leaf litter arthropod taxa in an
east Texas loblolly pine (Pinus taeda L.) plantation.
The objectives were (1) to measure the quantitative
response of this arthropod community to treatments of
different intensities and (2) to monitor the changes inthis arthropod community over a 2-year period
following tree harvesting and site preparation.
2. Methods and materials
2.1. Study site
The study site was located on land owned by
Temple-Inland Forest Products Corporation in Tyler
County, TX, USA. The site was located approximately
10 km south of Spurger, and just north of Fred, at
30.68N and 94.48W. Annual mean temperature is
19.4 8C and annual mean precipitation is 136 cm
(Griffiths and Bryan, 1987). Elevation ranges from 17
to 19 m above sea level.
The soil of the area is a BowieCaddoRains
association. This varies from an acid fine sandy loam
to an acid sandy clay loam, with variable drainage, and
a topsoil varying in depth from 12 to 45 cm. The
vegetation in the surrounding geographic area is
dominated by loblolly pine, longleaf pine (P. palustris
Mill.), shortleaf pine (P. echinata Mill.), and oaks(Quercus spp.).
Prior to harvesting, the site was a 30 ha, 27-year-
old loblolly pine stand established by direct seeding
and thinned in corridors at age 15 years. At least three
harvests of pine have occurred in the past from this site
and there is no history of cultivation. The stand was
partitioned into 28 plots. Each plot was approximately
28 m 42 m (0.12 ha) in size: large enough to allow a
14 14 grid of new seedlings to be planted on a 2 m
3 m spacing following tree harvesting and site
preparation. Four plots were allocated as a contiguousundisturbed reference area and situated approximately
200 m from the rest of the site.
2.2. Treatments
Two harvesting, two site preparation, and two
fertilization treatments were arranged in a 2 2 2
factorial to give eight different treatment combina-
tions in a randomized block design with three blocks
based on generalized soil texture and drainage. Hence,
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3 of the 24 plots were randomly allocated to each one
of these 8 combinations.
Trees were harvested either with a hand-fell, bole-
only-removal method or with a whole-tree mechanicalharvesting procedure. Hand-felling employed hand-
held chainsaws to harvest trees, the tree boles were
removed with cranes to minimize physical distur-
bance, and the foliage and branch material was left at
the site. The mechanical treatment involved harvesting
trees with a feller-buncher with rotary cutter and
skidders, and all foliage, branch, and bark material
was removed along with the tree bole. Harvesting was
conducted between July and August 1994, and a total
area of approximately 10 ha was harvested.
In September 1994, the harvested area of the site
was treated aerially with broadleaf herbicides (ima-
zapyr and triclopyr at 0.5 and 1.12 kg/ha, respec-
tively). In October 1994, half of the 24 harvested plots
were subjected to a bedding procedure in which
topsoil was arranged into elevated rows for seedling
planting. Rows were approximately 50 cm high and
50 cm wide separated by 50 cm wide furrows, and
coarse woody felling debris was removed from the
elevated surface. All beds appeared to be in good
condition for the duration of the study. Bedding was
applied in October 1994. Loblolly pine seedlings were
planted in March 1995, but due to pales weevil(Hylobius pales L.) damage, trees were pulled up, the
site re-treated with herbicide in September 1995, and
re-planted in February 1996. Harvested plots were
either left unfertilized or fertilizer was broadcast by
hand at a rate of 250 kg/ha with diammonium
phosphate (DAP) in May 1996. All treatments were
performed by Temple Inland Forest Products Corpora-
tion staff under supervision of R.N. Coulson.
2.3. Sampling
Core samples were taken using 10-cm deep, 5-cm
diameter plastic corers at random positions within
randomly selected plots. Individual sample points
were selected using random numbers. In bedded plots,
all samples were taken on the ridges of the elevated
beds and the depressions between beds were not
sampled. Cores were wrapped in aluminum foil and
packed in ice in thefield to immobilize arthropods and
to equilibrate litter and soil temperature (Mitchell,
1974). Sampling was done at approximately the same
time of day (between 11 a.m. and 3 p.m.) during each
visit to the site to minimize the effects of diurnal
fluctuations in abiotic conditions and the resulting
vertical movement of the arthropod fauna (Seastedtand Crossley, 1981). Cores were transported to a
laboratory and arthropods extracted using modified
Tullgren funnels. Arthropod samples were stored in
80% isopropyl alcohol. The fauna were identified to
suborder or family level, and abundance recorded to
generate an index of abundance for major taxonomic
groups.
Samples were removed from the site on a total of 24
dates between February 1994 and December 1996.
Five of those dates occurred before tree harvesting
occurred and 19 afterwards. Samples were taken at
approximately monthly intervals within this time
period. During the pre-harvest period, one core sample
was taken from four randomly selected plots of each
treatment block and two samples were taken from the
unharvested area. On each post-harvesting sampling
visit, two core samples were randomly taken from
plots of each of the eight treatment combinations and
two taken from the undisturbed plots.
2.4. Data analysis
Abundance data for numerically dominant arthro-pod taxa (Mesostigmata, Prostigmata, Oribatida, and
Collembola) were normalized with a log (x + 1)
transformation (Macfayden, 1962) and analyzed by
analysis of variance using StatViewTM software
(Abacus Concepts Inc., 1996). The DAP fertilization
treatment was not applied until May 1996, so data
were split into pre- and post-fertilization time periods
for analysis. Data from between February 1995 and
May 1996 (pre-fertilization) were analyzed with a
two-way ANOVA based on a randomized block with
three blocks and a 2
2 factorial assignment ofharvesting and site preparation treatments (Zar, 1996).
Data from between May to December 1996 (post-
fertilization) were analyzed with a three-way ANOVA
a s a 2 2 2 factorial. A critical level ofa = 0.05 was
used in all cases. Abundance data from the undis-
turbed plots were not included in the ANOVA due to
the lack of randomization and spatial independence of
these plots. Pre-harvest data also were omitted from
ANOVA due to imbalanced sampling effort across
treated plots.
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Mean abundance for different treatment combina-
tions, as well as for individual treatments, was
calculated from both pre- and post-fertilization time
periods to investigate any interaction between the
three treatment types (harvesting, bedding, and
fertilization). Total mean abundance of all arthropods
per core sample was calculated for each sampling date
to investigate temporal trends and overall response to
treatments.
3. Results
A diverse group of arthropod taxa was collected
from core samples. Acari (mites) were the most
abundant group collected, Oribatida being the
numerically dominant suborder in all samples. Data
analysis was focused on the four most abundant
arthropod groups: Mesostigmata, Prostigmata, Oriba-
tida, and Collembola.
3.1. Effects of harvesting technique
A varied response to harvesting treatment was
observed among arthropod taxa. During pre-fertiliza-
tion sampling, mean abundance of Mesostigmata,
Prostigmata, and Collembola was slightly higher in
treatment combinations that included hand-felling
compared to those that included mechanical-felling
(Figs. 14). During post-fertilization sampling, no
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208198
Fig. 1. Post-harvest mean abundance of Mesostigmata per coresample for pre-fertilization (February 1995April 1996) treatment
combinations. Error bars represent standard errors. Different letters
above bars indicate significant difference between treatments as
detected by ANOVA and Tukeys post-hoc testing.
Fig. 2. Post-harvest mean abundance of Prostigmata per core sample for pre-fertilization (February 1995April 1996) treatment combinations.
Error bars represent standard errors. Different letters above bars indicate significant difference between treatments as detected by ANOVA and
Tukeys post-hoc testing.
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consistent differences between the two harvesting
treatments were observed in Acari and Collembola
(Figs. 58). Mean abundance of Prostigmata, Astig-
mata, Oribatida, Protura, Collembola, Psocoptera, and
Isoptera per individual treatment (as opposed to
combinations of different treatment) was higher in the
hand-fell treatment (Tables 1 and 2). Araneae and
Pseudoscorpiones were not detected from mechani-cally-felled plots and detected in low abundance in
hand-felled plots (Table 1). ANOVA suggested no
significant differences in abundance of numerically-
dominant taxa between harvesting treatments prior to
fertilization (Table 2). Following fertilization, only
Collembola were significantly higher in hand-
felled plots, albeit at relatively low mean abundance
(Table 3).
3.2. Effects of bedding
During pre-fertilization sampling, Acari and
Collembola were higher in abundance in treatment
combinations that included the non-bedding treatment
compared to those that included bedding (Figs. 14).
These differences did not persist during post-
fertilization sampling, however (Figs. 58). Overall
mean abundance of Acari, Diplopoda, Protura,
Collembola, Isoptera, and Pselaphidae per individual
treatment was higher in the non-bedding treatment
(Table 1). Mean abundance of Symphyla, Psocoptera,
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208 199
Fig. 3. Post-harvest mean abundance of Oribatida per core sample
for pre-fertilization (February 1995April 1996) treatment combi-
nations. Error bars represent standard errors. Different letters above
bars indicate significant difference between treatments as detected
by ANOVA and Tukeys post-hoc testing.
Fig. 4. Post-harvest mean abundance of Collembola per core sample for pre-fertilization (February 1995April 1996) treatment combinations.
Error bars represent standard errors. Different letters above bars indicate significant difference between treatments as detected by ANOVA and
Tukeys post-hoc testing.
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and Formicidae were higher in the bedding treatment
(Table 1). Mesostigmata and Collembola were
significantly higher in abundance in non-bedded plots
during pre-fertilization sampling (Table 2), but no
significant post-fertilization differences were sug-
gested by ANOVA (Table 3). While not statisticallysignificant, mean abundance of Oribatida was
observed to be higher in bedded plots following
fertilization (Table 3).
3.3. Effects of fertilization
Fertilization had the most dramatic effect on
arthropod abundance. Mean abundance in treatment
combinations that included fertilization was higher for
all numerically dominant taxa (Figs. 58). Fertiliza-
tion led to an increase in the numerical dominance of
Oribatida, primarily at the expense of Collembola.
Overall mean abundance of Acari, Diplopoda,
Collembola, Isoptera, Psocoptera, and Formicidae
was higher in fertilized plots (Table 1). A few of the
rarer taxa, such as Diplura, Pselaphidae, and Dipteranlarvae, were lower in abundance in non-fertilized plots
(Table 1). ANOVA suggested significantly higher
abundance of Mesostigmata, Prostigmata, and Oriba-
tida following fertilization (Table 3).
3.4. Unharvested reference area
Arthropod abundance in the unharvested plots was
for the most part comparable to fertilized harvested
plots. However, Prostigmata were less abundant in the
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208200
Fig. 5. Post-harvest mean abundance of Mesostigmata per core sample for post-fertilization (MayDecember 1996) treatment combinations.
Error bars represent standard errors. Different letters above bars indicate significant difference between treatments as detected by ANOVA and
Tukeys post-hoc testing.
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harvested plots (Fig. 9) and Diplopoda, Isoptera, and
Staphylinidae were not detected from unharvested
plots (Table 4). Pauropoda was high in abundance and
Collembola and Psocoptera low in abundance
compared to treated plots (Fig. 9). Relative abundance
of Acari and Collembola was similar to treated plots(Tables 2 and 4; Fig. 9).
3.5. Temporal trend
Mean total abundance of all arthropods showed a
temporal trend of lower abundance in the hottest
months of the year (JuneSeptember) (Fig. 10). Mean
abundance was highly variable from month-to-month,
particularly following tree harvesting and site pre-
paration (Fig. 10). Highest abundance occurred
between November 1995 and March 1996. Total
abundance tended to be vary greatly between
individual samples on any one sample date.
4. Discussion
4.1. Overall disturbance
The cumulative disturbances caused by tree
harvesting and site preparation have many potential
abiotic and biotic effects on a forest ecosystem. With
removal of the tree canopy higher levels of solar
radiation reach the forest floor, organic matter input
patterns are altered, and temperature and moisture
fluctuations increase in the top 10 cm of the soil
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208 201
Fig. 6. Post-harvest mean abundance of Prostigmata per core sample for post-fertilization (MayDecember 1996) treatment combinations. Error
bars represent standard errors. Different letters above bars indicate significant difference between treatments as detected by ANOVA and Tukeys
post-hoc testing.
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(Weber and Methven, 1985; Bird and Chatarpaul,
1988). The mechanical tree felling and bedding
treatments used in this study have the potential to
alter nutrient dynamics and increase soil compaction,
soil erosion, and leaching losses (Tuttle et al., 1985).Due to these effects, clear-cutting and other silvicul-
tural practices have been recognized to have
significant effects on the invertebrate fauna of the
forestfloor (Heliovaara and Vaisanen, 1984; Hoekstra
et al., 1995). Despite these potential risks, in this study
the abundance of arthropods dwelling in the upper
10 cm of the forestfloor was not observed to decrease
significantly over a 2-year period following clear-
cutting. It should be noted, however, that 2 years is a
relatively short time period and longer term changes to
this arthropod community are feasible (Blair and
Crossley, 1988).
4.2. Tree harvesting and site preparation
Although only minimal pre-harvest data was
available for comparison, it appeared that there were
no overall and significant declines in total arthropod
abundance throughout the study above and beyond
expected seasonalfluctuations. Over 2 years following
tree harvesting, Acari, the numerically dominant taxa
collected from core samples, recovered rapidly in the
more intensive harvesting and site preparation
treatments. These trends suggest that the microenvir-
onmental conditions these arthropods were exposed to
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208202
Fig. 7. Post-harvest mean abundance of Oribatida per core sample for post-fertilization (MayDecember 1996) treatment combinations. Error
bars representstandard errors.Different letters above bars indicate significant difference between treatments as detected by ANOVA and Tukeys
post-hoc testing.
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stabilized during this time period, despite the removal
of tree canopy cover. The lack of significant
differences between harvesting and site preparation
treatments in 1996 suggests that the effects of
disturbance intensity caused by levels of each did
not persist during the course of the study. The lowermean abundance of some taxa, particularly Collem-
bola, in the mechanical-fell treatment could have
resulted from the removal of harvesting debris and the
more severe soil compaction caused by this treatment.
In mechanically-felled plots, where all felling debris
was removed, the soil surface was likely to be more
exposed to moisture and temperaturefluctuations and
available organic matter was likely reduced in
comparison to hand-felled plots. Collembola survival
and reproduction is strongly influenced by tempera-
ture and moisture and many species are detritivorous
(Christiansen, 1964; Huhta and Mikkonen, 1982),
which may explain the lower abundance observed in
the more intensively harvested plots.
The majority of similar studies have shown
significant reduction of Acari and Collembolaabundance over a number of years following tree
harvesting. Hence, the rapid recovery of Acari in this
study is surprising. Huhta et al. (1967, 1969) and
Huhta (1979) reported initial increases in mite and
springtail densities after clear-cutting followed by
significant declines. Other authors have reported
arthropod abundance decreases after tree harvesting
without initial increases (Vlug and Borden, 1973; Hill
et al., 1975; Lasebikan, 1975; Abbott et al., 1980;
Seastedt and Crossley, 1981; Bird and Chatarpaul,
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208 203
Fig. 8. Post-harvest mean abundance of Collembola per core sample for post-fertilization (MayDecember 1996) treatment combinations. Error
bars represent standard errors. Different letters above bars indicate significant difference between treatments as detected by ANOVA and Tukeys
post-hoc testing.
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1986; 1988; Cancela da Fonseca, 1990), including
impacts lasting up to 30 years (Blair and Crossley,
1988). Many Collembola are able to respond
numerically to disturbance with rapid reproduction
rates (Coleman and Crossley, 1996) so the lack of
significant recovery in this study could be seen as
surprising. Oribatida, however, tend to be slow
reproducers that take longer to recover from dis-
turbance so the abundance increase observed in this
study could also be viewed as unusual. Differences
between the findings reported here and other similar
studies may reflect geographic variation, the suit-
ability of specific silvicultural practices to different
forest types and locations, the size of study plot used in
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208204
Table 1
Mean abundance per core sample of selected arthropod taxa split by harvesting, site preparation, and fertilization treatments
Taxon Treatment
TH0 TH1 SP0 SP1 FE0 FE1
Araneae 0.08 0.00 0.05 0.03 0.00 0.11
Pseudoscorpiones 0.08 0.00 0.04 0.04 0.04 0.07
Acari 32.47 21.62 29.63 23.90 18.92 48.35
Chilopoda 0.05 0.07 0.08 0.04 0.04 0.04
Diplopoda 0.25 0.21 0.45 0.01 0.00 1.07
Pauropoda 0.01 0.04 0.03 0.03 0.04 0.07
Symphyla: Scolopendrellidae 0.32 0.33 0.25 0.40 0.50 0.32
Protura 2.36 1.03 2.07 1.33 1.39 1.82
Diplura: Japygidae 0.75 0.74 0.80 0.68 1.29 0.82
Collembola: Hypogastruridae 0.53 0.57 0.84 0.28 0.29 0.75
Isotomidae 7.11 4.24 5.82 4.16 3.21 3.86
Entomobryidae 1.79 1.13 2.16 0.76 0.46 1.29
Sminthuridae 1.18 0.86 1.33 0.72 0.25 0.29
Isoptera: Rhinotermitidae 0.21 0.05 0.24 0.03 0.07 0.61
Psocoptera 1.32 0.47 0.62 1.17 0.25 3.82
Thysanoptera: Phlaeothripidae 0.11 0.08 0.08 0.11 0.04 0.29
Coleoptera: Carabidae 0.00 0.11 0.07 0.04 0.00 0.04
Staphylinidae 0.04 0.09 0.07 0.07 0.04 0.04
Pselaphidae 1.17 1.00 1.37 0.79 1.21 0.82
Larvae 0.47 0.38 0.54 0.34 0.07 0.61
Diptera: Larvae 0.40 0.40 0.38 0.41 0.21 0.18
Hymenoptera: Formicidae 1.90 5.58 1.53 5.83 0.68 12.43
TH0, hand-fell, bole-only harvesting; TH1, mechanical-fell, whole-tree harvesting; SP0, non-bedding; SP1, bedding; FE0, non-fertilization;
and FE1, fertilization. Harvesting and bedding treatment means were calculated from a total of 228 samples taken from each treatment
between February 1995 and December 1996. Fertilization treatmentmeans were calculated from 126 samples taken between Mayand December
1996.
Table 2Mean abundance of soil and litter arthropods per core sample for the pre-fertilization sampling period (February, 1995 to May, 1996)
Treatment Mesostigmata Prostigmata Oribatida Collembola
Harvesting
Hand-fell, bole-only harvesting 4.6 (0.8)a 6.7 (1.6)a 44.5 (6.5)a 10.9 (1.7)a
Mechanical-fell, whole-tree harvesting 6.3 (2.2)a 5.6 (1.5)a 47.1 (10.4)a 9.0 (2.2)a
Bedding
Non-bedding 7.0(1.9)a 8.2 (2.0)a 50.4 (9.0)a 12.7 (1.9)a
Bedding 3.9(1.4)b 4.0 (0.8)a 41.3 (8.3)a 7.3 (2.0)b
Significant differences for each pair of treatments from two-way ANOVA are designated by different letters ( P < 0.05). Standard errors are in
parentheses. Data were transformed to log (x + 1) for all analyses. No significant interactions were detected between treatments.
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different studies, and the need for further research into
the responses of soil and litter arthropods to forest
management regimes.
The bedding treatment employed in this study
concentrates nutrients into upper soil layers for
utilization by newly planted tree seedlings. This
treatment also can increase the risk of wind and water
erosion and leaching losses. Nitrogen and phosphorus
levels have been shown to decrease following bedding
(Pye and Vitousek, 1985; Tew et al., 1986) and erosion
and soil compaction shown to increase (Tew et al.,
1986). The lower abundance of many arthropod taxa
observed prior to fertilization may have resulted from
these detrimental effects of bedding. The observation
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208 205
Table 3
Mean abundance of soil and litter arthropods per soil sample for the post-fertilization sampling period (May, 1996 to December, 1996)
Treatment Mesostigmata Prostigmat Oribatida Collembola
HarvestingHand-fell, bole-only harvesting 5.3 (1.5)a 12.8 (3.2)a 48.5 (14.9)a 6.8 (1.2)a
Mechanical-fell, whole-tree harvesting 4.8 (1.1)a 6.2 (1.4)a 39.5 (8.8)a 3.7 (0.8)b
Bedding
Non-bedding 6.8 (1.6)a 10.4 (2.4)a 36.0 (8.2)a 6.7 (1.3)a
Bedding 3.3 (0.8)a 8.6 (2.6)a 52.0 (15.1)a 3.8 (0.8)a
Fertilization
Non-fertilization 2.9 (0.6)a 4.8 (1.0)a 19.8 (3.5)a 4.3 (0.9)a
Fertilization 7.1 (1.7)b 14.2 (3.2)b 68.2 (15.7)b 6.3 (1.2)a
Significant differences for each pair of treatments from three-way ANOVA are designated by different letters (P< 0.05). Standard errors are in
parentheses. Data were transformed to log (x + 1) for all analyses. No significant interactions were detected between treatments.
Fig. 9. Comparison of mean abundance of selected arthropod taxa in post-harvest treated plots and unharvested plots. Error bars represent
standard errors.
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12/14
that post-fertilization abundance was comparable in
the two site preparation treatments for most taxaindicates that the arthropod community was suffi-
ciently intact to enable a recovery in bedded plots 2
years following tree harvesting.
Several taxa were of higher abundance in the
bedded plots than in non-bedded plots. Formicidae
were noticeably higher in mean abundance in both
bedded and mechanically-felled treatments.Solenop-
sis was the dominant genus among collected speci-
mens. These ants are, in general, omnivorous, highly
competitive, and successful at exploiting disturbed
habitats. Due to the eusocial nature of these insectshowever, spatial aggregation may cause these data to
be misleading when using the core sampling method.
4.3. Nitrogen and phosphorus fertilization
Varied responses of arthropod communities to
nitrogen and phosphorus fertilization have been
reported in other studies. For example, Kopeszki
(1993)concluded that the numerical response of Acari
and Collembola was determined by the mode of action
of the fertilizer used, whileHill et al. (1975) noted a
delayed effect of fertilization on arthropod abundancedue to a period of nutrient immobilization by
microorganisms. The immediate increase in arthropod
abundance following fertilization in this study
suggests that immobilization did not have a delaying
effect.
4.4. Arthropods, plant growth, and sustainable
timber production
Soil and litter arthropods have many interactions
with microorganisms in forest systems (e.g. seeWerner and Dindal, 1987) and positive feedbacks
exist between the activities of both. Considering these
relationships and the importance of many soil and
litter arthropods for decomposition and soil condition,
the increase in arthropod abundance following
fertilization has implications for future tree growth.
By regulating decomposition rates, organic matter and
soil aggregation dynamics, and soil aeration, the
recovery of the arthropod community observed here
could indicate a local environmental stabilization
S.B. Bird et al. / Forest Ecology and Management 202 (2004) 195208206
Fig. 10. Temporal trend in mean abundance of all soil and litter arthropods per core sample showing timing of treatment applications. Error bars
represent standard errors.
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7/25/2019 Changes in Soil and Litter Arthropod Abundance Following Three Harvesting and Site Preparation in a Loblolly Pine
13/14
leading to more optimal plant growth conditions.
These factors also have positive implications for
sustainability of timber production at this site when
using the combination of silvicultural practices
studied.
In conclusion, the overall rapid recovery of the
abundance of the soil and litter arthropod communityobserved in this study indicates that the silvicultural
practices used at this site may not have jeopardized
long-term site productivity. Arthropod diversity, soil
respiration, and soil nutrient data from the same site
support this conclusion (Messina personal commu-
nication, 1997;Bird et al., 2000). More extensive and
comparative studies of this nature are needed to
investigate the differences observed between different
geographic locations, forest types, and specific tree
harvesting and site preparation treatments.
Acknowledgments
We would like to thank M.G. Messina, M.C. Carter,
D.A. Crossley Jr., P.D. Teel, T.R. Seastedt, E. Rebek,M. Telfer, A. Gilogly, A. Bunting, J.E. Herrick, G.
Forbes, B. Sutter, A. Scott, and two anonymous
reviewers for manuscript review and helpful input. We
also thank P.E. Pulley and P.A. Dacin for statistical
advice. This study was made possible by a grant from
the Texas Research Enhancement Program and was
part of a collaborative study between the USDA Forest
Service, Temple Inland Forest Products Corp.,
Louisiana State University, and Texas A&M Uni-
versity.
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Taxon Mean abundance
Araneae 0.03
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