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Texas A&M University, Department of Ecosystem Science & Management, TAMU 2138, College Station, TX 77843, United States
USDA Forest Service, Agricultural Research Center, Normal, AL 35762, United States
a r t i c l e
i n f o
Article history:
Available online xxxx
Keywords:
Soil microbial biomass
Soil organic carbon
Soil total nitrogen
Forest harvest
Long-term soil productivity experiment
a b s t r a c t
Land management practices have strong potential to modify the biogeochemistry of forest soils, with
implications for the long-term sustainability and productivity of forestlands. The Long-Term Soil
Productivity (LTSP) program, a network of 62 sites across the USA and Canada, was initiated to address
concerns over possible losses of soil productivity due to soil disturbance from forest management.
Network sites employ an experimental design consisting of three harvest intensities (bole only, whole
tree, whole tree + forest oor removal) in combination with three soil compaction intensities (none,
intermediate, severe). Our purpose was to determine the impact of forest harvest intensity, soil compaction, and their interaction on soil organic carbon (SOC) and total nitrogen (TN) storage, and on soil
microbial biomass C and N (MBC and MBN, respectively) in a Pinus taeda L. forest 15-years post-treatment
at the Groveton LTSP site in eastern Texas, USA. Soils were sampled (010 cm) ve times during 2011
2012, and we quantied SOC and TN by dry combustion, and MBC and MBN by chloroform fumigation
extraction. SOC and TN were both higher in the bole only treatment compared to the more severe harvest
treatments; however, while TN was signicantly impacted by harvest and varied seasonally, SOC varied
only with season. MBC and MBN were impacted by harvest intensity and varied seasonally, and SMB-N
had a harvest by time interaction. Generally, both microbial indices decreased in the order: bole only
>whole tree > whole tree + forest oor. Temporal variations in MBN and TN were correlated with temperature. Soil compaction and the harvest intensity soil compaction interaction had no effect on the
measured soil properties. Since N limits tree growth in forest ecosystems, and because soil microbial biomass plays a key role in N mineralization, data suggest that harvest practices that minimize removal of
litter and slash will favor soil N retention, maintain the size of the soil microbial biomass pool, and maximize the potential productivity of future rotations.
2015 Elsevier B.V. All rights reserved.
1. Introduction
Forests in North America currently function as net carbon sinks
(Birdsey et al., 2007; Pacala et al., 2001); however, both natural and
anthropogenic disturbances can negatively impact the strength of
this sink (Chen et al., 2013; Dangal et al., 2014). Forest harvest
practices result in organic matter removal and may cause soil compaction, thereby enhancing the potential to impact forest productivity by altering the pool sizes of limiting nutrients and
inuencing rates of biogeochemical processes. When more than
the merchantable bole is harvested, increasing amounts of C and
Corresponding author at: University of Texas at San Antonio, Environmental
Science Programs, One UTSA Circle, San Antonio, TX 78249, United States. Tel.: +1
210 458 5798.
E-mail addresses: [email protected] (J.A. Foote), [email protected] (T.W.
Boutton), [email protected] (D.A. Scott).
nutrients such as N are exported off site in the non-bolewood tissues, potentially compromising soil fertility and the productivity of
subsequent forest rotations (Carter et al., 2002; Currie et al., 2003;
Henderson, 1995; Metz and Wells, 1965; Powers et al., 2005; Scott
et al., 2004; Turner and Lambert, 2011; Wells and Jorgensen, 1979;
Wollum and Davey, 1975). The legacy of these disturbance effects
on the carbon and nitrogen cycles of forest ecosystems may persist
for more than 50100 years (Chen et al., 2013; Kellman et al.,
2014). Additionally, soil compaction that occurs during forest harvest may increase bulk density and alter soil structure and porosity, which in turn decreases aeration, gas exchange, and water
inltration (Ampoorter et al., 2007; Berisso et al., 2012; Cambi
et al., 2015; Fisher and Binkley, 2000; Greacen and Sands, 1980;
Labelle and Jaeger, 2011; Powers et al., 2005), all of which can
inuence rates of biogeochemical processes in disturbed soils.
Declines in ecosystem C storage coupled with alterations in soil
http://dx.doi.org/10.1016/j.foreco.2015.03.036
0378-1127/ 2015 Elsevier B.V. All rights reserved.
Please cite this article in press as: Foote, J.A., et al. Soil C and N storage and microbial biomass in US southern pine forests: Inuence of forest management.
Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036
J.A. Foote et al. / Forest Ecology and Management xxx (2015) xxxxxx
The Gulf Coastal Plain region of the USA currently has some of
the highest rates of gross forest cover loss in the world (Hanson
et al., 2010). These rates are particularly high in the southwestern-most portion of this area where gross forest cover loss
exceeded 10% across large portions of this region between the
years 20002005 (Hanson et al., 2010). Despite the importance of
this land use activity, tree harvest effects on ecosystem biogeochemistry have been little studied in this region. Therefore, the
purpose of this study was to determine the impact of forest harvest
intensity, soil compaction and their interaction on SOC, TN, and soil
microbial biomass carbon and nitrogen (MBC and MBN, respectively) in a Pinus taeda L. (loblolly pine) forest 15 years posttreatment at the Long-Term Soil Productivity (LTSP) site located
near Groveton, Texas, USA. Three hypotheses were tested to
address the impact of forest harvest and soil disturbance on C and
N cycling: (1) SOC and TN will be lowest under the most severe tree
harvest and soil compaction treatments; (2) lower SOC and TN
under the most severe harvest treatments will constrain nutrient
availability to the current rotation, resulting in less aboveground
litter and more root biomass; and (3) greater losses of organic matter and soil structure changes due to compaction will result in
lower MBC and MBN in the most intense harvest treatments.
2. Materials and methods
2.1. Study area
Field sampling was conducted quarterly from March 2011
through March 2012 for a total of ve sample periods (March,
June, September and December 2011, and March 2012) at the
Long-Term Soil Productivity (LTSP) site in Davy Crockett National
Forest near Groveton, TX, USA (31060 32.4800 N, 95090 59.1500 W)
(hereinafter Groveton LTSP). The climate is subtropical with a
mean annual temperature of 19.1 C and mean annual precipitation of 1135 mm (19812010) that is bimodal, with peaks in
MayJune and October (Fig. 1). Topography is nearly at with
slopes of 13% and elevation ranging from 101 m to 110 m. Soils
across the study area are uniform (ne-loamy siliceous, thermic
Oxyaquic Glossudalf in the Kurth series) and developed in loamy
coastal plain sediments of the Yegua and Whitset geological formations. The A-horizon occurs at 015 cm, while the E-horizon
extends from 15 to 50 cm.
The Groveton LTSP treatment plots were established in 1997 in
accordance with the parameters specied by the LTSP program
(Powers, 2006) which consists of three harvest intensities (bole
only, whole tree, and whole tree + forest oor removal) and three
levels of soil compaction (none, intermediate, and severe) in
70
140
60
120
50
100
40
80
30
60
20
10
0
40
Temperature
Precipitation
n. b. ch ril ay e ly g. t. ct. v. c.
Ja Fe ar Ap M Jun Ju Au Sep O No De
M
20
Precipitation (mm)
physical structure that often follow tree harvest events may also
have adverse effects on the size of the soil microbial biomass pool,
and may also modify the structure and function of soil microbial
communities (Frey et al., 2009; Jordan et al., 2003; Vance and
Chapin, 2001; Wardle, 1992).
Forest harvest may result in the alteration of biogeochemical
cycling within the ecosystem; however, impacts are variable due
to differences in frequency, harvest method, climate, and inherent
site quality (Goetz et al., 2012). For example, a meta-analysis conducted by Johnson and Curtis (2001) found that harvest had no
overall impact on soil organic carbon (SOC) and soil total nitrogen
(TN). However, they did note that there were signicant differences between tree harvest methods. More specically, they
observed that SOC and TN were greater in bole only harvests and
lower in plots where the whole trees were harvested (Johnson
and Curtis, 2001). In contrast, a more recent meta-analysis by
Nave et al. (2010) found an overall decrease in forest soil C in
response to tree harvesting, but it was primarily related to the loss
of forest oor rather than mineral soil C-losses. Despite the generalizations that emerge from these two meta-analyses, individual
studies have shown that forest harvest intensity (i.e., removal of
more that the merchantable bole) may cause losses (Jones et al.,
2011; Kellman et al., 2014; Li et al., 2003;), gains (Grand and
Lavkulich, 2012; Vanguelova et al., 2010), or no change
(Jerabkova et al., 2011; Knoepp and Swank, 1997; Mariani et al.,
2006; McLaughlin and Phillips, 2006; Richter et al., 1999; Scott
et al., 2014; Wall, 2008; Zerpa et al., 2010) in SOC and TN.
It is unclear how soil compaction might inuence SOC and TN
storage and turnover. Compaction may destroy soil macroaggregates in the upper portion of the soil prole, thereby exposing soil
organic matter (SOM) to decomposers and accelerating decay processes (Six et al., 2004; Tisdall and Oades, 1982). Soil compaction
may also result in decreases in both root biomass and uptake of soil
mineral N, increasing the potential for ecosystem N losses via
leaching and denitrication (Jordan et al., 2003; Torbert and
Wood, 1992). Alternatively, compaction could restrict gas
exchange and reduce soil pore space, thereby limiting access of
decomposers to SOM resulting in no changes (de Neve and
Hofman, 2000; Cambi et al., 2015; Mariani et al., 2006; Sanchez
et al., 2006b) or possibly increases in SOC and TN (Tan et al., 2005).
Soil microorganisms are integral components of biogeochemical
cycles, and play key roles in the development and maintenance of
soil structure and fertility in forest ecosystems (Allen and
Schlesinger, 2004; Gallardo and Schlesinger, 1994; Wardle, 1992;
Zak et al., 1994). Soil microbial biomass as a living and active element of the soil may serve as a bellwether of changes in soil nutrient status resulting from management practices (Allen and
Schlesinger, 2004; Brookes, 2001; Haubensak et al., 2002;
Pregitzer, 2003; Wardle, 1992). Positive relationships have been
demonstrated between soil microbial biomass and SOC and TN
(Allen and Schlesinger, 2004; Brookes, 2001; Li et al., 2004;
Wardle, 1992); therefore, the removal or alteration of soil organic
matter inputs due to forest harvest may diminish the size of the
soil microbial biomass pool (Busse et al., 2006; LeDuc and
Rothstein, 2007; Li et al., 2004; Mummey et al., 2010; Tan et al.,
2005). However, other studies have found little effect of harvest
on soil microbial biomass (Busse et al., 2006; Mariani et al.,
2006). Soil compaction, by altering porosity, gas exchange, and
water inltration, has likewise resulted in variable impacts on soil
microbial biomass. Soil compaction may have no effect on soil
microbial biomass (Shestak and Busse, 2005; Busse et al., 2006),
lead to increases in soil microbial biomass due to reduced pore
space that limits predator access to microbes (Breland and
Hansen, 1996; Jensen et al., 1996a, 1996b; Li et al., 2004;
Mariani et al., 2006), or lead to decreases due to reduced aeration
and water conductivity (Frey et al., 2009; Tan et al., 2005).
Temperature ( oC)
Please cite this article in press as: Foote, J.A., et al. Soil C and N storage and microbial biomass in US southern pine forests: Inuence of forest management.
Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036
J.A. Foote et al. / Forest Ecology and Management xxx (2015) xxxxxx
Please cite this article in press as: Foote, J.A., et al. Soil C and N storage and microbial biomass in US southern pine forests: Inuence of forest management.
Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036
J.A. Foote et al. / Forest Ecology and Management xxx (2015) xxxxxx
temperature were conducted using the mean monthly temperature in the month prior to each sampling period. For each response
variable, correlations were run using the mean values for each plot
for every sample period, so that each correlation analysis was
based on 120 observations.
3. Results
Soil pH across the site was acidic and ranged from 3.5 to 4.7
(average 4.15 0.06, n = 27). Soil texture is loamy sand, with sand
concentration of 755 7 g kg1 and clay concentration of
65 3 g kg1 (n = 24).
Fig. 2. Weather conditions and soil moisture for January 2011 through April 2012.
(A) Crockett, Houston County, TX mean monthly temperature and total precipitation from January 2011 through April 2012, and (B) mean SE (n = 2427)
volumetric soil moisture for each sample period from 2011 to 2012.
Soil organic carbon content was not impacted by harvest intensity (p > 0.05; Table 2; Fig. 4A), but varied signicantly over time
(p < 0.05). These temporal variations were not correlated with air
temperature, precipitation, or soil moisture (Table 1). Average
Table 1
Pearsons correlation coefcients among variables examined in this study.
Precip.
Precipitation
Temperature
Volumetric soil moisture
Roots
Fine
Coarse
Total
Litter
SOC
TN
SOC/TN
Cmic/Corg
MBC
MBN
*
Temp.
0.48*
0.76*
0.71*
0.39
0.14
0.27
0.53*
0.01
0.12
0.13
0.09
0.26
0.17
0.04
0.35
0.34
0.04
0.25
0.32
0.11
0.04
0.22
0.53*
Soil moisture
0.56*
0.24
0.43
0.15
0.24
0.29
0.10
0.35
0.20
0.07
Roots
Fine
Coarse
Total
0.12
0.48*
0.15
0.23
0.04
0.39
0.21
0.48*
0.35
0.93*
0.17
0.02
0.05
0.09
0.13
0.12
0.06
0.22
0.10
0.07
0.23
0.18
0.28
0.06
Litter
SOC
TN
SOC/TN
Cmic/Corg
MBC
0.06
0.55*
0.67*
0.10
0.35
0.38
0.73*
0.36
0.59*
0.50*
0.44
0.38*
0.29
0.58*
0.64*
0.39
0.12
0.26
0.32
0.24
0.84*
MBN
p < 0.001.
p < 0.01.
p < 0.05.
Please cite this article in press as: Foote, J.A., et al. Soil C and N storage and microbial biomass in US southern pine forests: Inuence of forest management.
Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036
J.A. Foote et al. / Forest Ecology and Management xxx (2015) xxxxxx
Table 2
Results of repeated measures ANOVA (p-values) testing the effects of tree harvest
method, time, and their interaction on litter and root biomass, TN, MBN, SOC, MBC,
and Cmic/Corg.
Harvest
Time
Harvest time
0.384
0.942
0.782
0.793
0.055
0.004
0.368
0.166
0.892
0.668
0.479
0.581
0.425
0.048
0.032
0.006
0.988
0.028
0.003
0.013
0.004
0.012
0.100
0.215
0.299
0.047
0.255
p-Valuea
Litter
Fine roots
Coarse roots
Total roots
SOC (g C m2)
Total N (g N m2)
MBC (lg C g1)
MBN (lg N g1)
Cmic/Corg
a
p < 0.05;
p < 0.01;
4. Discussion
The removal of tree biomass coupled with soil disturbance during a forest harvest event can have lasting effects on both the quality and quantity of soil organic matter as well as soil physical
properties that can strongly control microbial activity and ecosystem biogeochemistry. Our results show that increased forest harvest intensity signicantly reduced soil TN and MBC 15 years
following harvest. Fine root mass, MBC, SOC, and TN varied over
the course of a single year, and there was a signicant harvest x
time interaction for MBN. However, neither soil compaction nor
its interaction with forest harvest intensity affected any of the
response variables we measured in this western Gulf Coastal
Plain site 15 years post-treatment.
p < 0.001
Please cite this article in press as: Foote, J.A., et al. Soil C and N storage and microbial biomass in US southern pine forests: Inuence of forest management.
Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036
J.A. Foote et al. / Forest Ecology and Management xxx (2015) xxxxxx
Fig. 3. Biomass (g m2) by time of sampling. (A) Litter; and (BD) roots. Symbols
are means SE (n = 3).
Please cite this article in press as: Foote, J.A., et al. Soil C and N storage and microbial biomass in US southern pine forests: Inuence of forest management.
Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036
J.A. Foote et al. / Forest Ecology and Management xxx (2015) xxxxxx
Fig. 4. (A and B) SOC and TN (g m2); (C and D) MBC and MBN (lg g1); and (E) Cmic/Corg by time of sampling. Symbols are means SE (n = 3). Cmic/Corg calculated using
concentrations of MBC and SOC.
very high soil moisture during March 2012 may have slowed the
recovery of the soil microbial biomass during that spring season.
The Cmic/Corg ratio varied signicantly over time, with higher
values during MarchJune 2011 compared to the September
2011March 2012 interval. Over the course of the entire study,
Cmic/Corg ratio was correlated negatively with SOC and the
SOC:TN ratio. Since Cmic/Corg is higher when SOC pool size is lower,
the microbial biomass is probably not carbon-limited. And, Cmic/
Corg is higher when the C/N ratio of the soil organic matter is lower.
Taken together, these observations suggest that the soil microbial
biomass pool in this study area is probably N-limited (Liao and
Boutton, 2008; Wardle, 1992).
4.4. Potential implications for forest productivity
Given that soil TN, MBC, and MBN were signicantly lower in
the more severe harvest treatments, is there any evidence that
these differences in key indices of soil fertility have affected tree
productivity at this site? At 5 years following the implementation
of treatments at the Groveton LTSP site, stand volumes were
6.88 m3 ha1 in the bole only, 5.13 m3 ha1 in the whole tree,
and 2.42 m3 ha1 in the whole tree + forest oor removal plots
Please cite this article in press as: Foote, J.A., et al. Soil C and N storage and microbial biomass in US southern pine forests: Inuence of forest management.
Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036
J.A. Foote et al. / Forest Ecology and Management xxx (2015) xxxxxx
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Forest Ecol. Manage. (2015), http://dx.doi.org/10.1016/j.foreco.2015.03.036