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2011 SOUTHEASTERN NATURALIST 10(3):489–500
Prescribed Fire and the Abundance of Soil
Microarthropods in Northeast Georgia
Matthew W. Hutchins¹,*, Barbara C. Reynolds¹, and Steven P. Patch²
Abstract - We examined the effects of prescribed fire on the abundance of soil microarthropods
in a southeastern pine-hardwood forest in northeast Georgia. Using soil cores,
the soil microarthropod groups Prostigmata, Oribatida, and Collembola were examined
before and after a low-intensity prescribed fire intended for fuels reduction and wildlife
habitat improvement. A post-burn evaluation found 100% duff layer coverage and 80%
of the understory vegetation consumed. Prostigmata numbers were significantly reduced
four months after the burn, with numbers returning to pre-burn levels more than one
year later. Although Oribatida and Collembola fluctuated from year to year, we found no
significant effects from the burn on those taxa. These results suggest that low-intensity
prescribed burning has no lasting negative effects on soil microarthropod populations.
The results from this study add to the evidence suggesting the adaptability of southeastern
forests to low-intensity prescribed fire.
Natural resource managers recognize fire as an important element in promoting
wildlife habitat, species composition, and protecting biodiversity
(Masters et al. 1996, Vose 2000). In addition to promoting habitat and species,
fire assists in reducing fuels along forested and urban interfaces. The use of
prescribed fire for stand maintenance and management is increasing in the
southeastern United States as managers observe benefits of fire and understand
the historical role that fire has played in southeastern ecosystems (Van
Lear 2000, Vose et al. 1999).
The southeastern mixed pine-hardwood ecosystem type is particularly dependent
on fire and may be negatively affected by fire suppression (Vose et al. 1997,
1999). Recent studies suggest low-intensity prescribed burning as an effective
method for restoring desirable species in southeastern pine-hardwood communities
without harmful effects on carbon and nutrient cycling (Hubbard et al. 2004,
Vose et al. 1999). However, even with desirable results of prescribed fire on nutrient
cycling, impacts on decomposer communities after fire could counterbalance
any positive results.
Prescribed burning alters crucial habitat and resources for soil decomposer
organisms through changes in above-ground vegetation, litter, and organic
matter (Haimi et al. 2000). With low-intensity prescribed burning, direct effects
to mobile soil invertebrates are of less concern than effects on immobile
¹Department of Environmental Studies, University of North Carolina at Asheville,
Asheville, NC 28804. ²Department of Mathematics, University of North Carolina at
Asheville, Asheville, NC 28804. *Corresponding author - email@example.com.
490 Southeastern Naturalist Vol. 10, No. 3
microorganisms. On the other hand, indirect effects of burning, such as litter
reduction, have shown to dramatically reduce abundance of mobile and immobile
soil-dwelling invertebrates (Certini 2005). Despite the impact fire has
on the forest floor habitat, soil fauna often display resilience to this type of
disturbance. Resilience may be defined not only as the stability of a system
following a disturbance, but how quickly the system is able to return to equilibrium
after disturbance (Holling 1973). The resilience of the soil system to
prescribed burning depends on factors such as climate, vegetation, and topography
of the burned area (Certini 2005). In addition, fire severity and depth of
burn has been suggested as the determining factor in the effects of fire on soil
fauna (Malmstrom 2010).
Microarthropods are among the most abundant organisms in forest soil and
serve vital functional roles in connecting the litter layer to mineral soil (Susilo
et al. 2004). Many microarthropods occupy multiple feeding guilds in the soil
food web as primary and secondary decomposers and feed on a range of materials
as shredders, scavengers, detritivores, and fungivores (Coleman et al. 2004,
Schneider et al. 2004). Other microarthropods are predators and feed on other
soil organisms, such as nematodes (Coleman et al. 2004). By breaking down organic
material, soil microarthropods increase the surface area of organic matter
for colonization of fungi and bacteria, indirectly impacting primary production
(Coleman et al. 2004, Crossley et al. 1992, Susilo et al. 2004). Fungal-feeding
microarthropods are also thought to be important in production of dissolved organic
matter (Osler and Sommerkorn 2007).
Soil mites and Collembola make up the majority of microarthropods
in most soil systems (Coleman et al. 2004). While the important role
of soil microarthropods is clear, there is still limited understanding of how the
soil microarthropod community is affected by prescribed burning or fire in
the southeastern US. The few investigations on this subject in the Southeast
are inconsistent in their findings. Studies on the impacts of fire on abundance
of mites, such as prostigmatids and oribatids, show variable results, with
some reports of lasting negative effects and other reports of no apparent effects
(Barratt et al. 2006, Coleman and Rieske 2006, Dress and Boerner 2004,
Haimi et al. 2000). A few studies indicate Collembola to be resilient to or
possibly benefit from disturbance, including fire (Coleman and Rieske 2006,
Lindberg and Bengtsson 2005), while other studies have implicated fire to
have lasting negative effects on abundance of Collembola (Berch et al. 2007,
Brand 2002, Collett 1998).
In this study, we examined the response of soil microarthropod populations
to a low-intensity prescribed burn in a southeastern pine-hardwood forest. Populations
of the mite suborders Prostigmata and Oribatida and the hexapod class
Collembola were examined at a burned treatment site, pre and post-burn, as well
as at an unburned control site.
2011 M.W. Hutchins, B.C. Reynolds, and S.P. Patch 491
The study area is located in northern Georgia, in the Lake Russell Wildlife
Management Area (USFS) in Habersham County. The area is about 480 m in
elevation, with a mean annual precipitation of 150 cm (NOAA 2007). The study
area consisted of two sites (burned and unburned) 0.4 km in distance from each
other. Both sites comprised slopes with north and south aspects. The unburned
south slope was 24°, the unburned north slope was 37°, the burned south slope
was 23° and the burned north slope was 27°.
Sites were mixed pine-hardwood forests, although the burned site consisted
of more pine in the overstory than the control site. The burned area was
predominantly Pinus taeda L. (Loblolly Pine) with some P. virginiana Miller
(Virginia Pine) in the overstory. Many pines at the burned site had recently died
due to pine bark beetle (Ips spp.) attacks, and the site had probably been tilled
for agriculture prior to forest growth (B. Boydstun, US Forest Service, Clarksville,
GA, pers. comm.). Understory trees at the burned site were mostly Acer
rubrum L. (Red Maple), Oxydendrum arboreum L. (Sourwood), and Nyssa sylvatica
Marsh. (Blackgum), with a few Cornus florida L. (Flowering Dogwood).
The unburned area was thought to be less disturbed even before the fire, with
forest that was previously harvested, but not used for agriculture (H.R. Pulliam,
University of Georgia, Athens, GA, pers. comm.). The overstory at the unburned
site was comprised of Virginia Pine, Quercus prinus L. (Chestnut Oak),
Q. alba L. (White Oak), and Carya spp. (Hickory), with some Q. falcata Michx.
(Southern Red Oak). The understory at the unburned site consisted of Red
Maple, Flowering Dogwood, Sourwood, Blackgum, and Southern Red Oak,
with some Kalmia latifolia L. (Mountain Laurel) on the north slope. Young Red
Maple saplings were the predominant understory on the south slope at the unburned
Site treatment and sampling
In April 2005, one site was burned by the USDA Forest Service using a mix
of hand-set and helicopter-set ignitions on a grid basis as part of a prescribed
fire that burned 328 ha. The main purpose of the prescribed fire was fuels reduction
along a wildland/urban interface. Secondary benefits of the prescribed fire
included wildlife habitat improvement. Surface air temperatures on the day of
the burn ranged from 16–18 °C when the burn started in the morning to 24 °C by
the afternoon. Relative humidity ranged from 35–40% when the burn started to
about 27% by the afternoon. The intensity of the fire was primarily influenced by
temperature and relative humidity on that day. The site was burned in the morning
at approximately 11:00 am EST; therefore, the intensity of the burn was considered
light to moderate due to the temperature and relative humidity at the time
of the burn (B. Boydstun, pers. comm.). A post-burn evaluation of the area after
the fire found no bare soil exposed, with 100% of the duff layer (layer between
492 Southeastern Naturalist Vol. 10, No. 3
soil and leaf litter) coverage and 80% of the understory vegetation consumed (B.
Boydstun, pers. comm.).
For our study, a 200-m² area was divided into ten 1- x 2-m plots on both
north and south slopes at each site, totaling 40 plots among both burned
and unburned sites. North and south plot areas were about 50 m apart at the
burned site and about 80 m apart at the unburned site. Plots were divided into
an imaginary grid of 12 cells, and collections were made from a randomly
selected cell within the grid, using the same cell on a given date. Every plot
was sampled on each collection date, for a total of 20 samples/site. Soil cores
were taken with PVC piping, 5 x 5 cm (98 cm³) with 1.5-mm mesh screen on
one end, through the litter layer (Moldenke 1994). Pre-burn soil core samples
were taken on 9 July 2004. Post-burn soil core samples were taken on 20 August
2005 and on 12 September 2006, four and 17 months after the burn. Both
burned and unburned sites were sampled each sampling date and the unburned
site was considered the control. Soil temperatures were also taken at each
plot with soil thermometers to depths of 3–5 cm, and soil moisture was measured
using a Hydrosense© (Campbell Scientific, Inc.) water content sensor at
depths of 12 cm.
The soil cores were wrapped in aluminum foil and transported in a cooler back
to the lab the same day and immediately placed screen-side down on modified
Tullgren extractors (Mallow and Crossley 1984). Microarthropods were collected
into vials containing 70% ethanol over one week with the light intensity gradually
increased. Soil microarthropods were sorted into the categories Prostigmata, Oribatida,
and Collembola. Other microinvertebrate taxa, such as pseudoscorpions,
were separated but not analyzed in this investigation due to their low numbers. To
investigate the effects of fire on oribatid life history, we separated the oribatids
into two groups: mature and immature. Immature oribatids were considered to
be mites that were smaller (<1 μm) and that had weakly sclerotized cuticles. We
recognize that many immature oribatids are endophagous, and thus would not
respond to the Tullgren extraction (Norton 1994). Therefore, our counts of immature
oribatids are probably low.
A linear mixed model was used to analyze the data for each microarthropod
taxa: prostigmatids, immature oribatids, mature oribatids, and Collembola. This
analysis assumes that the burned treatment site and the unburned control site had
the same properties relative to the effects of fire on microarthropods. Because
there was only one burn, it was not possible to test this assumption. However,
the two sites were adjacent to each other and there were no obvious differences
in the geographical features between the two sites. The responses were the natural
log counts of taxa. A natural log transform was used to make the residuals
homoskedastic and normally distributed (Littell et al. 2006). Taxa count data did
not contain excess zeros, thus, there was no need to accommodate zero inflation
(Sileshi 2008). Plot was considered as a random factor. Treatment (burned and
2011 M.W. Hutchins, B.C. Reynolds, and S.P. Patch 493
unburned) and aspects (north and south) were between-plot factors; sampling
date and the interaction between sampling date and treatment were within-plot
factors. For each microarthropod taxon, plots of the residual versus fitted values
were consistent with homoskedastic residuals. There was no evidence of nonnormality
in the residuals for any taxon (all Shapiro-Wilks P-values > 0.10).
Because the natural log transformed counts were used, inferences were made for
the geometric mean counts instead of the mean counts.
Least square means and standard errors were used for comparison purposes of
dates and any significant interactions using the family significance level of 0.05
with Tukey adjustment for pair-wise comparisons. The resulting 95% confidence
intervals for the means of the log transformed counts were reverse transformed
to obtain corresponding confidence intervals for geometric mean counts. These
confidence intervals for geometric mean counts had individual confidence levels
A total of 6685 microarthropods were extracted from our soil cores, including
810 prostigmatid mites (10% of total), 3259 immature oribatids (50% of total),
1830 mature oribatids (28% of total), and 786 Collembola (12% of total). The
abundance of immature oribatids were mainly attributed to the increased abundance
one year after the burn in summer 2006 (Table 1).
All four microarthropod taxa (prostigmatids, immature oribatids, mature
oribatids, and Collembola) were most abundant in the summer of 2006, one
year following the burn. Prostigmatid populations differed by date and had
a significant treatment*date interaction (F = 5.42, P = 0.0063). Prostigmata
abundance was lower at the burned treatment site in 2005 compared to 2004,
but returned to pre-burn levels in 2006 (Fig. 1). The treatment*date interaction
for Prostigmata presumably indicates a significant effect from the burn.
Within the oribatids, both immature and mature groups were significantly
more abundant by date with no treatment interaction; thus, fluctuations in
populations were similar in plots at both burned and unburned sites. Immature
oribatids were significantly more abundant in summer 2006 than summer 2004
Table 1. Abundance of immature oribatids, mature oribatids, and Collembola. Numbers are geometric
means (GM) of all burned and unburned samples and represent 95% confidence intervals
(CI) for geometric mean counts. For each taxon, means with the same letter are not significantly
Pre-burn Four month post-burn 17 month post-burn
(Summer 2004) (Summer 2005) (Summer 2006)
Avg. # / soil Avg. # / soil Avg. # / soil
Taxon core (GM) 95% CI core (GM) 95% CI core (GM) 95% CI
Immature oribatids 7.72 (B) 5.05–11.56 4.36 (B) 2.74–6.68 36.48 (A) 25.15–52.71
Mature oribatids 12.39 (A) 9.08–16.78 3.92 (B) 2.72–5.51 15.23 (A) 11.27–20.47
Collembola 3.80 (B) 2.76–5.13 3.11 (B) 2.23–4.24 7.29 (A) 5.51–9.56
494 Southeastern Naturalist Vol. 10, No. 3
Figure 2. Collembola at burned treatment and unburned control sites. Bars are means
combining 2004, 2005, and 2006 data at each site and represent individual 95%
confidence intervals for geometric mean counts. Bars with the same letter are not significantly
Figure 1. Effect of prescribed burning on Prostigmata. Bars are means at the burned treatment
and unburned control sites and represent individual 95% confidence intervals for
geometric mean counts. Bars with the same letter are not significantly different.
2011 M.W. Hutchins, B.C. Reynolds, and S.P. Patch 495
or summer 2005 (F = 35.56, P < 0.0001; Table 1). However, unlike the immatures,
mature oribatid abundance decreased from 2004 to 2005, and then increased
from 2005 to 2006 (F = 23.35, P < 0.0001; Table 1). The abundances
of mature oribatids were similar in 2004 and 2006. Collembola abundance
differed between treatment and control sites with no interaction with date or
aspect (F = 7.46, P = 0.0097). Collembola also differed by date with no interaction
with treatment or aspect (F = 9.56, P = 0.0002). Collembola were more
abundant at the treatment site, regardless of year, pre and post-burn (Fig. 2).
Among sampling years, Collembola were also more abundant in summer 2006
than in summer 2004 and summer 2005 (Table 1). Aspect was not significant
for any microarthropod group.
The low-intensity prescribed fire had negative effects on the abundance
of prostigmatid mites four months after the burn in 2005, but their populations
returned to pre-burn levels 17 months post-burn. While the populations
of each microarthropod taxon fluctuated over the sampling period, oribatids
and Collembola showed no effects from the burn. Barratt et al. (2006) and
Seastedt (1984) found Prostigmata to be unaffected by fire, but others found
lasting negative effects of prescribed burning on microarthropod taxa including
Prostigmata (Barratt et al. 2006, Berch et al. 2007, Cole et al. 2008, Dress
and Boerner 2004). Also, Coleman and Rieske (2006) examined impacts of
prescribed burning on ground-dwelling and leaf-litter arthropods in a southeastern
forest and found differential effects among the two groups. While
leaf-litter arthropods were negatively affected for two years following a
prescribed fire, ground-dwelling arthropods were apparently unaffected (Coleman
and Rieske 2006).
Negative short-term effects of fire on microarthropods have been shown to
depend on the intensity and frequency of burning (Barratt et al. 2006, Dress
and Boerner 2004). Fire disturbs nutrient cycling and biotic components of the
ecosystem, but also provides sources of nutrients and resources to the soil food
web in the form of charcoal and burned vegetation. This reciprocal effect of fire
disturbance results in direct and indirect impacts to soil microarthropods (Hart
et al. 2005). Fire directly affects microarthropods by causing mortality due to
temperature or combustion. Temperature thresholds, related to intensity and
duration of fire, have been demonstrated as direct effects to soil microarthopods
(Haimi et al. 2000, Malmstrom 2008). While some microarthropods dwell deeply
enough in the soil horizon to be insulated from even high-severity fire (Coleman
and Rieske 2006, DeBano et al. 1998), the samples from our study were not deep
enough to include these organisms. Indirect effects of fire on microarthropods
include changes in habitat (resulting in increased vulnerability to environmental
variables) and resources (such as food sources) (Barratt et al. 2006, Brand 2002,
496 Southeastern Naturalist Vol. 10, No. 3
Most studies attribute the effects of fire on forest-dwelling arthropods to the
the amount and quality of leaf litter remaining after a fire-disturbance event
(Coleman and Rieske 2006, Donegan et al. 2001, Dress and Boerner 2004).
Generally, Prostigmata, Oribatida, and Collembola are positively correlated with
organic matter (Peterson and Luxton 1982, Vreeken-Bruijs et al. 1997), although
one study has found exceptions to this assumption (Hasegawa 2001). Organic
matter in the litter layer influences microarthropod abundance by creating nutrient
pools for soil microarthropod food sources, such as the microbial community
(Neary et al. 1999). Considering that most Prostigmata are predators and depend
on prey (Coleman and Crossley 2004), perhaps our observation of their decrease
in 2005 was the bottom-up effect of a decrease in their prey (other microinvertebrates)
that rely on microbial activity.
Oribatids were not affected by the burn, but they have been reported to be
suppressed by a wide range of disturbances due to their slow reproduction rates
(Cancela da Fonseca and Sarkar 1998, Cole at al. 2008, Crossley et al. 1992,
Lindberg and Bengtsson 2005). We found different responses between the immature
and mature oribatids over the three years of the experiment. Immature
oribatid abundance was relatively low in 2004 and 2005, with a large increase
in 2006, seeming to indicate fluctuation between years. The numbers of mature
oribatids significantly fluctuated year to year during the sampling period, but
again, changes in abundance were apparently unrelated to the burn as fluctuations
occurred at both sites.
Collembola significantly increased from 2005 to 2006 at both the burned
treatment site and unburned control site. Collembola were also more abundant
at the burned treatment site compared to the control including pre-burn, with no
interaction with year, perhaps reflecting site differences, such as land-use history
and current canopy cover. The higher predominance of pine at the burn site also
may have attributed to site differences. Collembola have been shown to respond
positively to disturbance events (Coleman et al. 2004). Other studies have found
that Collembola respond negatively to fire (Berch et al. 2007, Brand 2002, Collett
1998). However, these studies tended to examine effects of fire on species richness
and diversity, which we did not address.
While our study did not measure nitrogen in the litter-layer, the presence
of increased soil-litter nitrogen following the burn may have contributed to
the stability of oribatids and Collembola. Nitrogen is often the most limiting
nutrient in ecosystems, but soil nitrogen often increases directly after a
low-intensity fire (DeBano et al. 1998, Elliot et al. 2004). Nitrogen is known
to stimulate the growth of fungi, an important food source for Oribatida and
Collembola (Vreeken-Buijs et al. 1997). Perhaps an initial increase of nitrogen
following the burn gave a temporary boost to Oribatida and Collembola
populations in the burned treatment site, superseding any negative effects of
organic matter or litter loss.
Because there was only one fire disturbance, our findings depend on the
assumption that the disturbance location and control location had the same
2011 M.W. Hutchins, B.C. Reynolds, and S.P. Patch 497
properties relative to the effects of fire on soil microarthropods. While the preburn
data helps support the sampling of the disturbance event, the effects, or lack
of effects, found in this study could be partially due to differences in site location.
Ideally, a study would have several small fire disturbances created in random locations,
which would allow one to adjust tests of disturbance effects for variation
in the implementation of creating the fire disturbance and the properties of the
locations of fire disturbance, but as with many disturbance studies, that was not
possible in this setting.
Our results assist in understanding the effects of single low-intensity burning
in southeastern pine-hardwood forests. Our findings suggest Prostigmata
abundance was significantly decreased by the low-intensity prescribed burn, but
recovered one year following the burn. This short-term negative response by
the Prostigmata was most likely due to temporary loss of habitat and resources.
The other microarthropod taxa—immature oribatids, mature oribatids, and
Collembola—were apparently unaffected by the prescribed burn, which probably
demonstrates their opportunistic feeding behavior. The results from this study
also indicate the great extent to which environmental variables affect the abundance
of microarthropod populations. Our study demonstrates the need to further
understand the underlying controls on microarthropod abundance and the links
among organic matter, microbial activity, and taxa of soil microarthropods. Because
soil microarthropods are indicators of soil quality and serve important roles
in the soil food web, our results should assist resource managers in understanding
some ecological implications of prescribed burning in the southeastern US. Our
findings add to the literature suggesting that soil systems in the Southeast are
highly adaptable to fire disturbances.
We would like to give a special thanks to Irene Rossell, Ernest C. Bernard, and two
anonymous reviewers for their valuable comments and suggestions. We would also like
to thank Blaine Boydstun (USFS), Stephanie Madson, H. Ronald Pulliam, and Scott Eustice
for their help. We also gratefully acknowledge NSF LTER grant NSF-DEB 0218001,
which supported this study.
Barratt, B.I., P.A. Tozer, R.L. Wiedemer, C.M. Ferguson, and P.D. Johnstone. 2006.
Effect of fire on microarthropods in New Zealand indigenous grassland. Rangeland
Ecology and Management 59:383–391.
Berch, S.M., J.P. Battigelli, and G.D. Hope. 2007. Responses of soil mesofauna communities
and oribatid mite species to site-preparation treatments in high-elevation
cutblocks in southern British Columbia. Pedobiologia 51:23–32.
Brand, R.H. 2002. The effect of prescribed burning on epigeic springtails (Insecta: Collembola)
of woodland litter. American Midland Naturalist 148:383–393.
498 Southeastern Naturalist Vol. 10, No. 3
Certini, G. 2005. Effects of fire on properties of forest soils: A review. Oecologia
Cancela da Fonseca, J.P., and S. Sarkar. 1998. Soil microarthropods in two different managed
ecological systems (Tripura, India). Applied Soil Ecology 9:105–107.
Cole, L., S.M. Buckland, and R.D. Bardgett. 2008. Influence of disturbance and nitrogen
addition on plant and soil animal diversity in grassland. Soil Biology and Biochemistry
Coleman, D.C., Crossley, D.A., Jr., and P.F. Hendrix. 2004. Fundamentals of Soil Ecology.
Second Edition. Elsevier Academic Press, Burlington, MA. 386 pp.
Coleman, T.W., and L.K. Rieske. 2006. Arthropod response to prescription burning at the
soil-litter interface in oak-pine forests. Forest Ecology and Management 233:52–60.
Collett, N.G. 1998. Effects of two short-rotation prescribed fires in autumn on surfaceactive
arthropods in dry sclerophyll eucalypt forest of west-central Victoria. Forest
Ecology and Management 107:253–273.
Crossley, D.A., B.R. Mueller, and J.C. Perdue. 1992. Biodiversity of microarthropods in
agricultural soils: Relations to processes. Agriculture, Ecosystems, and Environment
DeBano, L.F., D.G. Neary, and P.F. Ffolliott. 1998. Fire’s Effects on Ecosystems. John
Wiley and Sons, Inc. New York, NY. 333 pp.
Donegan, K.K., L.S. Watrud, R.J. Seidler, S.P. Maggard, T. Shiroyama, L.A. Porteous,
and G. DiGovanni. 2001. Soil and litter organisms in Pacific Northwest forests under
different management practices. Applied Soil Ecology 18:159–175.
Dress, W.J., and R.E. Boerner. 2004. Patterns of microarthropod abundance in oakhickory
forest ecosystems in relation to prescribed fire and landscape position. Pedobiologia
Elliot, K.J., J.M. Vose, B.D. Clinton, and J.D. Knoepp. 2004. Effects of understory
burning in a mesic mixed-oak forest of the southern Appalachians. Tall Timbers Fire
Ecology Conference Proceedings 22:272–283.
Haimi, J., H. Fritze, and P. Moilanen. 2000. Responses of soil decomposer animals to
wood-ash fertilization and burning in a coniferous forest stand. Forest Ecology and
Hart, S.C., T.H. DeLuca, G.S. Newman, M. Derek, and S.I. Boyle. 2005. Post-fire vegetative
dynamics as drivers of microbial community structure and function in forest
soils. Forest Ecology and Management 220:166–184.
Hasegawa, M. 2001. The relationship between the organic matter composition of a forest
floor and the structure of a soil arthropod community. European Journal of Soil
Holling, C. 1973. Resilience and stability of ecological systems. Annual Review of Ecology
and Systematics 4:1–23.
Hubbard, R.M., J.M. Vose, B.D. Clinton, K.J. Elliott, and J.D. Knoepp. 2004. Standrestoration
burning in oak-pine forests in the southern Appalachians: Effects on
aboveground biomass and carbon and nitrogen cycling. Forest Ecology and Management
Lindberg, N., and J. Bengtsson. 2005. Population responses of oribatid mites and collembolans
after drought. Applied Soil Ecology 28:163–174.
2011 M.W. Hutchins, B.C. Reynolds, and S.P. Patch 499
Littell, R.C., G.A. Milliken, W.W. Stroup, R.D. Wolfinger, and O. Schabenberger. 2006.
SAS® for Mixed Models. Second Edition. SAS Institute Inc. Cary, NC.
Mallow, D., and D.A. Crossley, Jr. 1984. Evaluation of five techniques for recovering
postlarval stages of chiggers (Acarine: Trombiculidae) from soil habitats. Journal of
Economic Entomology 77:281–284.
Malmstrom, A. 2008. Temperature tolerance in soil microarthropods: Simulation of
forest-fire heating in the laboratory. Pedobiologia 51:419–426.
Malmstrom, A. 2010. The importance of measuring fire severity: Evidence from microarthropod
studies. Forest Ecology and Management 260:62–70.
Masters, R.E., C.W. Wilson, G.A. Bukenhofer, and M.E. Payton. 1996. Effects of pinegrassland
restoration for Red-cockaded Woodpeckers on White-tailed Deer forage
production. Wildlife Society Bulletin 24:77–84.
Moldenke, A.R. 1994. Arthropods. Pp. 517–542, In R.W. Weaver, S. Angle, P. Bottomley,
D. Bezdicek, S. Smith, A. Tabatabai, and A. Wollum (Eds.). Methods of Soil Analysis.
Part 2: Microbiological and Biochemical Properties. Soil Science Society of America,
Inc., Madison, WI. 1692 pp.
National Oceanic and Atmospheric Association (NOAA). 2007. NOAA website. Available
online at http://www.noaa.gov. Accessed 9 May 2007.
Neary, D.G., C.C. Klopatek, L.F. DeBano, and P.F. Ffolliott. 1999. Fire effects on belowground
sustainability: A review and synthesis. Forest Ecology and Management
Norton, R.A. 1994. Evolutionary aspects of oribatid mites life histories and consequences
for the origin of the Astigmata. Pp. 99–135, In M.A. Houck (Ed.). Mites: Ecological
and Evolutionary Analyses of Life-History Patterns. Chapman and Hall, New York,
NY. 357 pp.
Osler, G.H.R., and M. Sommerkorn. 2007. Toward a complete soil C and N cycle: Incorporating
the soil fauna. Ecology 88:1611–1621.
Peterson H., and M. Luxton. 1982. A comparative analysis of soil fauna populations and
their role in decomposition processes. Oikos 39:287–388.
SAS Institute, Inc. 2003. SAS version 9.1 for Windows. Cary, NC.
Seastedt, T. 1984. Microarthropods of burned and unburned tallgrass prairie. Journal of
the Kansas Entomological Society 57:468–476.
Schneider, K., S. Migge, R.A. Norton, S. Scheu, R. Langel, A. Reineking, and M.
Maraun. 2004. Trophic niche differentiation in soil microarthropods (Oribatida, Acari):
Evidence from stable-isotope ratios (15N/14N). Soil Biology and Biochemistry
Sileshi, G. 2008. The excess-zero problem in soil animal count data and choice of appropriate
model for statistical reference. Pedobiologia 52:1–17.
Susilo, F.X., A.M. Neutel, M. van Noordwijk, K. Hairiah, G. Brown, and M.J. Swift.
2004. Pp. 285–307, Soil biodiversity and food webs. In M. van Noordwijk, G. Cadisch,
and C.K. Ong (Eds). Below-ground Interactions in Tropical Agroecosystems.
CABI Publishing Cambridge, MA. 440 pp.
Van Lear, D. 2000. Recent advances in the silvicultural use of prescribed fire. Tall Timbers
Fire Ecology Conference Proceedings 21:183–189.
Vose, J.M. 2000. Perspectives on using prescribed fire to achieve desired ecosystem conditions.
Tall Timbers Fire Ecology Conference Proceedings 21:12–17.
500 Southeastern Naturalist Vol. 10, No. 3
Vose, J.M., W.T. Swank, B.D. Clinton, R.L. Hendrick, and A.E. Major. 1997. Using fire
to restore pine/hardwood ecosystems in the southern Appalachians of North Carolina.
Pp. 149–154, In Fire Effects on Rare and Endangered Species and Habitats Conference
Proceedings, November 13–15, 1995. International Association of Wildland
Fire, Fairfield, WA. 343 pp.
Vose, J.M., W.T. Swank, B.D. Clinton, J.D. Knoepp, and L.W. Swift. 1999. Using standreplacement
fires to restore southern Appalachian pine-hardwood ecosystems: Effects
on mass, carbon, and nutrient pools. Forest Ecology and Management 114:215–226.
Vreeken-Buijs, M.J., J. Hassink, and L. Brussaard. 1997. Relationships of soil microarthropod
biomass with organic matter and pore-size distribution in soils under different
land use. Soil Biology and Biochemistry 30:97–106.