Open Journal of Forestry
2012. Vol.2, No.3, 97-109
Published Online July 2012 in SciRes (http://www.SciRP.org/journal/ojf) http://dx.doi.org/10.4236/ojf.2012.23013
Herbaceous-Layer Community Dynamics along a
Harvest-Intensity Gradient after 50 Years of Consistent
Management
Marcella A. Campione*, Linda M. Nagel, Christopher R. Webster
School of Forest Resources and Environm en tal Science, Ecosystem Science Center,
Michigan Technological University, Houghton, USA
Received April 16th, 2012; revised May 18th, 2012; accepted June 10th, 2012
In 1958, a demonstrational cutting trial totaling 22.2 ha was established in a northern hardwood forest in
Alberta, MI. Eight different treatments were installed, including four diameter-limit tre atments (56 cm, 41
cm, 30 cm, and 13 cm), three single-tree selection treatments with residual basal areas of 21 m2·ha–1, 16
m2·ha–1, and 11 m2·ha–1, and an uncut control. Within each treatment, a 0.4-ha permanent plot was estab-
lished and subdivided into 0.04-ha square subplots. Harvests have been implemented every ten years with
the most recent harvest occurring during the winter of 2008-2009. We quantified ground layer vegetation
response before and after the most recent harvest. Nonmetric multidimensional scaling (NMS) ordination
showed a very distinct separation between the most intensive management treatment (13-cm diame-
ter-limit treatment) and the uncut control. Compositionally, the diameter-limit treatments moved with
greater directionality and magnitude towards the 13-cm diameter-limit treatment following harvest, while
compositional change in the residual basal area treatments was less pronounced and lacked strong direc-
tionality. Herbaceous species percent cover generally decreased with increasing residual overstory basal
area across treatments. Weedy and early successional species were most abundant under lower residual
basal area and diameter-limit treatments. Results based on 50 years of continuous management suggest
that diameter-limit harvests likely have a greater impact on the herbaceous community than single-tree
selection or no management.
Keywords: Northern Hardwood Forests; Uneven-Aged Management; NMS Ordination; Understory
Diversity; Diameter-Limit; Single-Tree Selection
Introduction
While overstory dynamics in forested ecosystems have re-
ceived considerable study (e.g. Nyland, 1996; Oliver & Larson,
1996; Frelich, 2002), herbaceous-layer dynamics in response to
natural and anthropogenic disturbance are less well understood,
especially given the contribution of this layer to biological di-
versity and ecosystem function (Gilliam, 2003). Within north-
ern temperate forests in North America, the herbaceous layer
often contains the highest species richness (Curtis, 1959; Whit-
taker, 1967; Whitney & Foster, 1988; Scheiner & Istock, 1994;
Gilliam, 2007) and represents a disproportionate amount of the
net primary productivity relative to its biomass (DeAngelis et
al., 1981).
Land-use often leaves a legacy across a landscape that pro-
vides an important historical context for interpreting contem-
porary vegetation dynamics. The Great Lakes region was once
dominated by a vast forest; the United States General Land
Office Survey estimated circa 1850 that there were approxi-
mately 32.6 million ha of closed-canopy forests with 47% or
15.3 million ha in the hardwood forest type (Frelich, 2002).
Widespread logging in the Great Lakes region began in the
mid-1800s, with an estimated 20 million ha of forested land
harvested in 60 years (Williams, 1989). The rapid pace of
commodity driven harvesting and associated slash fires left
millions of hectares of cutover and degraded forests (Williams,
1989; Stearns, 1997).
Contemporary forest management in northern hardwood for-
ests in the Great Lakes region has focused on producing sus-
tainable, high quality sawtimber using uneven-aged regenera-
tion harvest methods (e.g., Arbogast, 1953; Arbogast, 1957;
Tubbs, 1977) with forest diversity as a lower priority. However,
decreases in understory tree diversity have been a common con-
sequence of long-term, uneven-aged management in northern
hardwood forests (Leak & Sendak, 2002; Kelty et al., 2003;
Neuendorff et al., 2007; Gronewold et al., 2010). Studies in
northern hardwood forests have shown varying responses in the
herbaceous layer due to uneven-aged management (e.g. Metz-
ger & Schultz, 1981; Crow et al., 2002; Scheller & Mladenoff,
2002; Kern et al., 2006; Wolf et al., 2008; Burton et al., 2009).
Along the border of northern Wisconsin and the Upper Pen-
insula of Michigan, Scheller and Mladenoff (2002) observed
that actively managed uneven-aged northern hardwood stands
had significantly greater herbaceous species richness than
old-growth northern hardwood stands. Uneven-aged stands
received management approximately ten to thirteen years prior
to sampling. The greatest percent cover in all herbaceous spe-
cies groups (ferns, forbs, weeds, graminoids , and shrubs) except
*Present Address: Department of Wildland Resources, Utah State Univer-
sity, Logan, UT 84322, U S A.

M. A. CAMPIONE ET AL.
spring ephemerals was observed in the managed uneven-aged
stands. Spring ephemerals may be more sensitive to uneven-
aged management due to reoccurring disturbance on the forest
floor and in the canopy (Metzger & Schultz, 1981; Scheller &
Mladenoff, 2002). However, at the Argonne Experimental For-
est in northern Wisconsin, there was no observed difference in
spring or summer herbaceous species between areas receiving
no management, uneven-aged, or even-aged management nine
years after the most recent harvest (Kern et al., 2006). This lack
of species composition change between management types (un-
even- or even-aged management) was also observed in a central
Indiana hardwood study (Jenkins & Parker, 1999).
Land-use history, including timber harvest, is a factor that
can affect herbaceous species composition that is often not
controlled for in studies, or is unknown (e.g. Scheller & Mlad-
enoff, 2002). In addition, time since harvest may be important
in assessing vegetation dynamics. For example, when harvest
intensities were controlled in the Argonne Experimental Forest,
pre-harvest conditions were not sampled and post-harvest con-
ditions were measured a minimum of nine years after harvest
activities (Kern et al., 2006). The lack of pre-harvest data and
the delay in sampling could have missed important composi-
tional changes in vegetation that occurred before and after har-
vest. These compositional changes may be transient; however,
repeated short cutting cycles in northern hardwood forests have
been hypothesized to create novel herbaceous communities
dominated by transient early successional species (Scheller &
Mladenoff, 2002).
Few long-term studies of the impact of forest management
on the herbaceous layer in northern temperate forests are avail-
able (e.g. Kern et al., 2006). One of the longest running and
most consistently treated silvicultural trials in the upper Great
Lakes region began in 1957 as a demonstrational woodlot at the
Ford Forest (Michigan Technological University, Alberta, MI)
with the goal of assisting small landowners by providing exam-
ples of various management techniques for northern hardwood
forests (Bourdo & Johnson, 1957; Reed et al., 1986; Erickson et
al., 1990; Bodine, 2000). The consistency of the study offers a
unique opportunity to observe the response of the herbaceous
layer after 50 years of management. The overstory and under-
story of the study site are dominated by Acer saccharum, and
the herbaceous layer adds the majority of species richness and
diversity in these stands. Our primary objective was to observe
how herbaceous-layer plant communities that have developed
under various management approaches respond to contempo-
rary harvesting. We hypothesized that herbaceous species com-
position and response to harvesting would vary along a gradient
of harvest intensity, and that the less intensively managed treat-
ment would have herbaceous communities more similar to the
second-growth, uncut control than the more intensively man-
aged treatments.
Methods
Site Description an d S tudy History
The study is located at the Ford Forest, owned by Michigan
Technological University and located in Alberta, Michigan
(46.66˚N, 88.51˚W; Baraga County). The site once supported a
Pinus strobus-hardwood forest with the pine resource being
removed around 1890 (Bourdo & Johnson, 1957). By 1938, the
Ford Motor Company had “selectively logged” the study site
and the surrounding area twice before the area was donated to
Michigan Technological University in 1954 (Bourdo & John-
son, 1957). Further information about pre-treatment conditions
and harvest guidelines can be found in Bourdo and Johnson
(1957). The demonstrational woodlot reached its fifth cutting
cycle and 50th year of harvest activity during the winter of
2008-2009. Consistent management has occurred over this
period with strict adherence to a 10-year cutting cycle (Bourdo
& Johnson, 1957).
The site’s proximity to Lake Superior regulates temperatures;
17.4˚C and –9.8˚C are the average summer and winter tem-
peratures, respectively (Berndt, 1988). Average total precipita-
tion and snowfall are 87.4 cm and 385.5 cm, respectively
(Berndt, 1988). The soils of the area are classified as Allouez
gravelly coarse sandy loams with slopes generally ranging from
0% - 6% (Berndt, 1988). The original composition of the over-
story was mainly Acer saccharum, with Tilia americana,
Betula alleghaniensis, and Ulmus americana as important as-
sociate species. Acer saccharum has increased in dominance in
all layers of vegetation with uneven-aged management (Reed et
al., 1986; Erickson et al., 1990; Bodine, 2000).
Study Design an d D at a C ol l ecti on
The study consists of eight different harvest treatments rang-
ing in size from 1.2 to 5.7 ha with a total study area of 22.2 ha.
There are four diameter-limit treatments of 56 cm (22 in), 41
cm (16 in), 30 cm (12 in), and 13 cm (5 in); three single-tree
selection treatments with residual basal areas of 21 m2·ha–1 (90
ft2·ac–1), 16 m2·ha–1 (70 ft2·ac–1), and 11 m2·ha–1 (50 ft2·ac–1);
and an uncut control. Diameter-limit treatments are defined as
the removal of any tree of any species over the specified di-
ameter (Helms, 1998). The residual basal area treatments gen-
erally focus on the removal of poor quality trees in all size
classes (12.7 cm and greater diameter at breast height (1.37 m))
with managemen t generally following a q-factor of 1.3 (Schwartz
et al., 2005). The maximum residual diameter for trees was set
at 61 cm (24 in). The uncut control has not received active
management since 1938 with the exception of a sanitation cut
to remove Ulmus americana in the 1980’s. The first harvest
occurred during the winter of 1958-1959 when all treatments
were harvested. The 21 m2·ha–1 residual basal area treatment,
the 41 cm diameter limit treatment, the 16 m2·ha–1 residual
basal area treatment, and 11 m2·ha–1 residual basal area treat-
ment have been harvested during the last five cutting cycles
(cutting cycle: 10 yrs). The 56-cm diameter-limit treatment,
30-cm diameter-limit treatment, and the 13-cm diameter-limit
treatment were cut in four, three, and two of the last five cutting
cycles, respectively.
The treatments are not replicated and generally follow a gra-
dient of harvest intensity beginning with the 21 m2 ha-1 residual
basal area treatment to the 13-cm diameter-limit treatment (Fig-
ure 1). Within each treatment a 0.4-ha permanent block was
established by Bourdo and Johnson in 1957 (Figure 1). The
permanent block was subdivided into ten 0.04-ha square sub-
plots. Within each subplot, all overstory species (greater than
11.4 cm in dbh) were identified to species and measured for
diameter, height, tree grade, and number of 2.4-m logs or 2.4-m
sticks.
During the 1998 pre-harvest sampling, seedling and sapling
plots were established in each subplot to quantify the estab-
lishment and recruitment of different tree species (Bodine,
98

M. A. CAMPIONE ET AL.
Compartment Treatment
021 Uncut Control
003 21 m2·h a–1 (90 ft2·ac–1)
004 56 cm DLH (22 in)
005 16 m2·h a–1 (70 ft2·ac–1)
006 41 cm DLH (16 in)
007 11 m2·ha–1 (50 ft2·a c–1)
008 30 cm DLH (12 in)
009 13 cm DLH (5 in)
Figure 1.
Ford Forest (Michigan Technological University) silviculture cutting
trial designed by Eric Bourdo in 1957. The study totals 22.2 ha which is
subdivided into nine different treatments. Within each treatment, there
is a 0.4-ha permanent rectangular block which is subdivided into ten
0.04-ha subplots. Within each subplot, one 0.008-ha circular plot used
to measure saplings was located in the center. Three 0.0004-ha circular
plots used to measure seedlings were set equidistance from the center.
At the center of each 0.0004-ha circular plot, a 1-m2 quadrat was used
to estimate herbaceous percent cover. All plot locations are perma-
nently marked. DLH refers to diameter limit harvest.
2000). A 0.008-ha circular plot was established at the center of
each subplot to measure sapling density, totaling 10 plots per
treatment. Sapling classification was based on both height and
diameter: 1) 30.5 cm to 91.4 cm in height, 2) 91.5 cm in height
to 2.41 cm dbh, 3) 2.43 cm to 7.49 cm dbh, and 4) 7.5 cm to
11.42 cm dbh. Three circular 0.0004-ha plots were located
equidistant from the center of the subplot where seedling den-
sity was measured, totaling 30 plots per treatment (Figure 1).
Seedlings were defined as any woody tree individual less than
30.5 cm height. During the 2008 pre-harvest sampling, addi-
tional 1-m2 quadrats were established to measure the impact
each treatment was having on the diversity and composition of
the herbaceous layer. At the center of the seedling plots, a 1-m2
quadrat was placed facing north to estimate percent coverage of
herbaceous species, totaling 30 plots per treatment (Figure 1).
Percent cover was estimated for each species; total percent
cover in each quadrat could total more than 100%. Herbaceous
species were identified to species except when accurate identi-
fication was not possible in the field; in those cases, species
were identified to genus. Overstory sampling occurred during
July and August of 2008 and 2009. Sampling of all layers of
understory vegetation occurred during July in both 2008 and
2009 (pre- and post-harvest, respectively). Data collected from
2008 and 2009 was used for analysis. Lack of pretreatment
herbaceous vegetation data (1957) limits our ability to make
inferences related to change over time. However, the habitat
type is consistent across the treatments (ATD, Acer-Tsuga-
Dryopteris; Burger and Kotar 2003) with similar species com-
position across all treatments.
Data Analysis
Nonmetric multidimensional scaling (NMS) ordination was
used to examine herbaceous species composition along a gra-
dient of harvesting intensity. NMS was used due to the relaxed
assumption of normality and because it does not assume a lin-
ear response in species to different gradients (McCune & Grace,
2002). Data were organized at the subplot level; each treatment
contained 10 subplots for a combined total of 80 subplots for
each year, 2008 (pre-harvest) and 2009 (post-harvest) sampling
periods (n = 160). Unknown species were deleted prior to
analysis; unknown species were present in only two of the 160
plots used in the ordination. The remaining herbaceous species
percent cover data were square-root transformed, which is a
common transformation used for percent cover data to reduce
the influence of a few samples with high percent cover (Field et
al., 1982). Transformed herbaceous species percent cover by
treatment and year was arranged in n-dimensional space using
PC-Ord Version 5 (McCune & Mefford, 2011). Autopilot mode
(slow and thorough) was selected using the Sørensen (Bray-
Curtis) distance measurement and a random starting configura-
tion. Two hundred and fifty runs were completed for both the
real data and randomized data to determine dimensionality.
Correlation analysis in the statistical interface R (R Develop-
ment Core Team, 2009) was used to test environmental vari-
ables used in the ordination for significance.
Summary statistics for each treatment were calculated at the
subplot level (n = 10) for overstory tree basal area per hectare,
herbaceous percent cover, and seedling and sapling stems per
hectare. Species richness (S) and Shannon’s diversity index (H’)
were calculated for the herbaceous layer, seedling, and sapling
layers. The herbaceous species richness is the average number
of species per 1-m2 quadrat.
Results
Overstory Composition
The 13-cm diameter-limit harvest was the only treatment not
harvested during the winter of 2008-2009; this treatment was
last harvested during the winter of 1998-1999. All other ac-
tively managed treatments were harvested with the specifica-
tions established by Bourdo and Johnson in 1957. As expected,
management decreased overstory basal area in all treatments
receiving active management (Figure 2). The average diameter
at breast height (dbh) of trees harvested in the permanent plots
of each diameter-limit treatment was 55.4 cm (st.dev = 0.5) for
the 56-cm diameter-limit treatment, 31.2 cm (st.dev = 12.8) for
the 41-cm diameter-limit treatment, and 31.5 cm (st.dev = 7.3)
for the 30-cm diameter-limit treatment. In the 41-cm and 30-cm
diameter-limit treatments, a few trees under the minimum di-
ameter were removed due to operability constraints, but there
was no tending in the smaller diameter classes. The residual
basal area treatments typically removed smaller diameter trees;
the average dbh of removed trees was 20.4 cm (st.dev = 10.7)
for the 21 m2·ha–1 residual basal area, 20.2 cm (st.dev = 7.9) for
the 16 m2·ha–1 residual basal area, and 24.4 cm (st.dev = 11.5)
for the 11 m2·ha–1 residual basal area. The diameter-limit treat-
ments generally created larger openings across the treatments,
resulting in higher light environments (personal observation).
Herbaceous Species Composition
Fifty-two herbaceous species were observed during both pre-
and post-harvest sampling periods. Twelve exotic species were
observed, representing 23% of the total species richness (Table
1). Slightly more species were observed in the post-harvest A

M. A. CAMPIONE ET AL.
100
21 m
2
·ha
(90 ft
2
·ac) 56 cm DLH
(22 in) 16 m
2
·ha
(70 ft
2
·ac) 11 m
2
·ha
(50 ft
2
·ac)
41 cm DLH
(16 in) 30 cm DLH
(12 in) 13 cm DLH
(5 in)
Control
White = 2008
Gray = 2009
52%
0%
29.1%
13.9%
6.9%
7.0%
8.2%
0%
45
40
35
30
25
20
15
10
5
0
Basal Area (sq·m/hectare)
Figure 2.
Overstory basal area per hectare pre- and post-harvest summarized by subplot (n = 10) at the Ford Forest Cut-
ting Trial, Alberta, MI. Percentage above each treatment represents the average decrease in basal area (m2·ha–1).
DLH refers to diameter limit harvest.
Herbace ous an d Woo dy Rich ness (S ) and Di versi ty (H’ )
period (39 observed in 2009 versus 35 observed in 2008). Of
these 52 species, nine species only occurred once during the
two years. On average more herbaceous species were observed in each
treatment in 2009 than in 2008 except in the 16 m2·ha–1 residual
basal area treatment and the 13-cm diameter-limit treatment
(Table 1). There was an average of 8.5 herbaceous species
observed in the 13-cm diameter-limit treatment in 2008 and an
average of 9.6 herbaceous species observed in the 11 m2·ha–1
residual basal area treatment in 2009 (Table 2). However, di-
versity of herbaceous species on average decreased from 2008
to 2009 (Table 2).
Total percent cover of the herbaceous layer in all treatments
increased from 2008 to 2009; the greatest increase occurred in
the 13-cm diameter-limit treatment, increasing from an average
of 15.5% in 2008 to 105.4% in 2009 (Table 1). Dryopteris
spinulosa, Carex spp., Rubus spp., and Galeopsis tetrahit were
herbaceous and semi-woody species that showed the greatest
increase in average percent cover from 2008 to 2009 (Table 2).
Dryopteris spinulosa consistently increased in percent cover
from 2008 to 2009 in all treatments; Rubus spp. also increased
in all treatments except for the uncut control and the 30-cm
diameter-limit treatment (Table 2). The largest average percent
cover increases for Dryopteris spinulosa occurred in the uncut
control, 1.9% to 10.3% respectively (Table 2). Rubus spp. had
the largest average percent cover increase in the 13-cm diame-
ter-limit treatment, 6.9% to 56.1% respectively (Table 2). Dry-
opteris spinulosa and Rubus spp. had high frequencies across
all treatments an d between years (Table A2).
The control contained on average the fewest number of spe-
cies in each of the understory layers in 2008 and 2009, except
for the sapling layer in 2008 (Table 2). The 21 m2·ha–1 residual
basal area treatment had on average 2.2 species of seedlings
observed (Table 2). Diversity was generally low in all layers of
the understory for the control (Table 2).
Compositional Change in the Herbaceous Layer
The NMS ordination solution was three-dimensional, ex-
plaining 84% of the variance in herbaceous community compo-
sition, and had a final stress of 13.94 (Figures 3 and 4). Axis 2
and Axis 3 were the most informative axes, explaining 37% and
26% of the variation, respectively. Axis 2 and Axis 3 were
significantly associated with distance to stream (m), overstory
basal area, average diameter of overstory, and sapling Shan-
non’s diversity (Table 3). Total seedling density and sapling
richness were also significantly associated with Axis 3 (Table
3). Total sapling density, seedling richness, seedling Shannon’s
diversity, overstory richness, overstory Shannon’s diversity,
percent of Acer saccharum seedling, sapling, and overstory
layers, and distance from the road are additional variables that
were included in the analysis, but did not have a significant
ffect.
Few species were found in only one treatment which may be
due to the proximity of treatments to one another (Table A3;
Figure 1). Herbaceous species co-occurrence in the actively
managed treatments generally ranged between 40-60% during
both years (Table A3). The uncut control and the 13-cm di-
ameter-limit treatment became more dissimilar from 2008 to
2009; a 32% herbaceous species overlap was observed in 2008
vs. 10% herbaceous species overlap in 2009 (Table A3). The
uncut control and the 13-cm diameter-limit harvest share few
species in common. In 2009, only Dryopteris spinulosa and
Trillium spp. were observed in both treatments. The 30-cm
diameter-limit treatment had the greatest herbaceous species
overlap with the 13-cm diameter-limit treatment in 2008 and
2009, 52% and 46% respectively (Table A3). e

M. A. CAMPIONE ET AL.
Table 1.
Herbaceous and woody species abundance, richness (S), and Shannon’s Diversity Index (H’) pre- and post-harvest by treatment at the Ford Forest
ial, Alberta, MI. Means are ± one standard deviation in parentheses. Seedlings are defined as any woody tree species less than 30.5 cm in
rol (90 ft2·ac–1) (22 in) (70 ft2·ac–1)(16 in) (50 ft2·ac–1) (12 in) (5 in)
Cutting Tr
height. Saplings are defined as any woody tree species greater than 30.5 cm in height and less than 11.42 cm at dbh. Herbaceous species richness is
the average number of species per 1-m2 quadrat. Herbaceous percent cover, seedling, and sapling stem/hectare are summarized by subplot (n = 10).
DLH refers to diameter limit harvest.
Cont 21 m2·ha–1 56 cm DLH16 m2·ha–1 41 cm DLH11 m2·ha–1 30 cm DLH 13 cm DLH
208 0
Herbaceous Percent Cover 2.) 7.)6.)3.) 15.5)
1.(1.) 3.) 4.(1.)5.(1.)7.(1.)8.(2.) 6.) 8.(1.)
Seedlins/ha)
Sapli/ha)
Herbacet Cover
(1.) 4.) 6.(2.)5.(2.)8.(2.)9.(3.) 7.) 6.(1.)
Seedlins/ha) 5
Sapli/ha)
8 (1.96.1 (2.6) 1(2.84.7 (12.2)3(2.98.6(2.4) 7 (1.45(1.
SHerb 8 19 (0.9906498343 (2.55 2
H’Herb 0.42 (0.40) 1.06 (0.25) 1.28(0.19)1.49(0.3)1.88(0.21)1.77(0.36) 1.7 (0.42) 1.57(0.18)
g (stem1067 (436) 517 (387) 187(179)304(160)304(210)11(5) 272 (142) 57 (39)
SSeedling 1.4 (0.5) 1.8 (0.4) 1.9(0.3)2.2(0.9)2.7(0.7)2.2(0.9) 3.1 (0.3) 2.1 (0.7)
H’Seedling 0.03 (0.05) 0.2 (0.26) 0.4(0.26)0.3(0.3)0.57 (0.34) 0.45(0.3) 0.93 (0.18) 0.55(0.34)
ng (stems110 (54) 46 (31) 97(60)96(58)96(55) 112 (44) 59 (17) 22 (13)
SSapling 2.4 (1) 2.2 (1.0) 3.2(0.6)2.5(1.1)3 (0.5)3.2(0.6) 3.2 (0.4) 2.7 (0.7)
H’Sapling 0.21 (0.21) 0.43 (0.32) 0.68(0.29)0.45(0.27)0.44(0.12)0.63(0.29) 0.7 (0.32) 0.71(0.36)
2009
ous Percen14.7 (9.8) 30.3 (25.5) 47.5(27.9)17.7(17.6)19.9(15)41.5(21.4) 21.9 (15.7) 105.4(26.7)
SHerb 2 28 (1.8134116627 (2.47 3
H’Herb 0.35 (0.36) 0.89 (0.41) 1.05(0.39)1.08(0.41)1.29(0.42)1.37(0.49) 1.27 (0.42) 1.03(0.19)
g (stem46(280) 291 (214) 96(63) 165(138)190 (97)97(52) 77 (61) 666(524)
SSeedling 1.2 (0.4) 3 (1.1) 3.1(1.0)2.9(1.3)3.7(1.0)3.4(1.0) 3.2 (1.0) 1.3 (1.0)
H’Seedling 0.02 (0.04) 0.5 (0.27) 0.86 (0.29) 0.62 (0.44) 0.89 (0.38) 0.93 (0.33) 0.95 (0.28 0.28(0.39)
ng (stems271 (157) 42 (29) 91(58)164(104) 97(56) 101 (38) 43 (19) 32 (16)
SSapling 3.1 (1.5) 3.3 (1.4) 4.6(1.1)4 (0.9)4.1(1.1)4.3(1.3) 4 (1.3) 3.8 (1.2)
H’Sapling 0.15 (0.13) 0.66 (0.26) 0.91(0.35)0.55(0.3)0.58(0.24)0.8(0.33) 0.93 (0.37) 0.93(0.2)
Table 2.
Mean percent cover (± one standard deviation in parentheses) of selectederbaceous species pre- and post-harvest at the Ford Forest Cutting Trial,
. All herbaceous species with average percent cover greater th 1% are included. Percent cover is summarized by subplot (n = 10). Full
(90 ft·ac) (22 in) 16 m·ha (70 ft2·ac–1) 41 cm DLH (16 in) 11 m·ha (50 ft2·ac–1) 30 cm DLH (12 in) 13 cm DLH (5 in)
h
anAlberta, MI
species names are in Table A1. DLH refers to diameter limit harvest.
Species Code Control 21 m2·ha–1
2–1 56 cm DLH 2–1 2–1
2008
carspp 1.1 (0.4)4 (0.4)
1.) 1.) 1.(0.) 1(0.)
2.3 (0.3)
1.9 (0.7) 6.9 (0.4)
ad 3.(3. )
1.(1. )1.(0.) 3.) 3(4.)
1.3 (1.3) 1.5 (0.8)
10.3 (3.0) 7.1 (2.0) 8.9 (2.3)1.5 (0.5)
2.3 (0.5)
4.3 (1.3)
1.3 (1.3)
1.7 (1.3) 1.5 (0.8) 6.3 (2.7) 2.3 (1.3)
4.1 (1.7)
1.9 (1.9)
20.5 (5.5) 25.7 (6.4)4 (2.5)5.3(3.1)13.7(4.4)
drycar 9 (0.56 (0.36 4.13
galtet*
rubspp 3.2 (0.7) 2.9 (0.5)
2009
iped 4 2
carspp 5 2 6 79 (1.84.98
cautha
drycar 6.2(2.5)3.7(1.4)1.4 (0.8) 1.3 (1.3)
elyhys
galtet* 1.2(0.6)2.2(1.0)14.6(4.9) 8.6 (3.4) 4.4 (3.6)
lapcom*
loncan
oryasp
polspp
ribspp 1.3 (1.2) 2.4 (2.3)
rubspp 56.1(5.1)
*Exoti c species.

M. A. CAMPIONE ET AL.
Axis 2
Axis 3
2
2
–2
–2
–0.8
–0.4 0.4
0.8
0.0 0.0
Saplings
Richness
Saplings
Shannon’s
Diversity
Distance to
Stream (m)
Total
Seedlings
Average
Diamrter o
f
Overstroy
Overstroy
Basal Area
adiped
dicspp
diovil
myospp
geuspp
galspp
actspp
mairac
linbor ribspp
drycar
aritri
sancan
alltri
triaur
marsa
trispp
osmchi cautha
viospp
carpen ortasp
ortasp
santri
osmc la
cirlut
lapcom
veroff
loncan
polspp
rubspp ipospp
carspp
aranud
eurspp
taroff
vertha
fraspp
leuvul
staspp
galtet
gaupro
antcot el yhys
hieaur
helthel
samspp
maican
menarv
rhacat
jefdip
braerec
Figure 3.
Non metric multi dimensional scaling ordination of herbaceous species observed in each treatment in 2008 and 2009 at the Ford Forest Cut-
I. Axis 2 explains 37% of the variation while Axis 3 explains 26% of the variation. All species bolded with asterisks are
T
se
f
Discussion
After 50 years of m observed two distinct
herbaceous communitiecontrol and the 13-cm
the
years, which may be due to
di
ting Trial, Alberta, M
exotic. The solid black ellipsis is associated with the 13-cm DLH. The black dotted ellipsis is associated with the Control. Full species names
and additional information can be found in Table A1. The insert is the significant environmental variables (p = 0.05) and their relation to or-
dination space. Additional information about significant environmental variables can be found in Table 3.
he uncut control and the 13-cm diameter-limit harvest were
d to compare how the actively managed treatments shifted diameter-limit treatments. There was some movement in herbaceous community between u
ater harvest. These two treatments were used due to the dis-
tinct communities that have developed in the last 50 years of
management in the 13-cm diameter-limit treatment and pre-
sumably due to the lack of active management in the control
(Figures 4(a) and (b)).
The diameter-limit treatments generally shifted composition-
ally with greater directionality and magnitude towards the
13-cm diameter-limit treatment compared to the residual basal
area treatments (Figures 4(a)-(e)). Compositional movement in
the residual basal area treatments was generally smaller in
magnitude and was also more random in direction (Figures
4(a), (b), (f) and (g)).
anagement, we
s in the uncut
fferences in environmental conditions; no harvesting occurred
in either of these treatments bet ween 2008 and 2009. The NOAA
National Weather Service station at Alberta, MI recorded lower
average temperatures during the months of June and July in
2009 and higher precipitation falling in the form of snow and
rain during the months of April, May, and June in 2009 prior to
sampling. This increased precipitation and cooler conditions
may be one of the main reasons why percent cover of herba-
ceous species increased in all treatments. In treatments where
harvests did occur, increased light levels may have also con-
tributed to increases in percent cover in the herb layer. This
movement could also be attributable to the carousel effect; the
basic premise of which is that individual species in a plant
community are not spatially static through time but rather tend
to reoccur in similar locations (van der Maarel and Sykes,
1993). In a closed canopy forest in Stockholm, Sweden under-
story species composition changed little within the forest be-
102

M. A. CAMPIONE ET AL.
Axis 2
Axis 3
2
2
1 1
Axis 3
2
–2
–2
–1
–1 1–2
Axis 2
2
–1
–1
–2
1
0
0
0
0
(a) (b)
Axis 2
Axis 3
2
2
–2
–2
1
–1 1
22
11
–1
Axis 2
Axis 3
Axis 3
2
–2
–2 –1 1
Axis 2
2
–2
–2 –1 1
–1
–1
(c) (d) (e)
Axis 3
2
2
–2
–2
1
–1 1
–1
Axis 2
Axis 3
2
2
–2
–2
1
–1 1
–1
Axis 2
Axis 3
2
2
–2
–2
1
–1 1
–1
Axis 2
(f) (g) (h)
Figure 4.
Non metric multi dime Forest Cutting Trial,
Alberta, MI: (a) 1 3-c m (f) 11 m2·ha–1 residual;
a–1 residual; and (h) 21 m2·ha–1 residual. Axis 2 explains 37% of the variation while Axis 3 explains 26% of the variation. Dots
nsional scaling ordination of pre- and post-harvest conditions for each treatment at the Ford
DLH (diameter-li mit harv est); (b ) Co ntrol; (c) 3 0-c m DLH; (d ) 41 -cm DLH; (e) 56 -cm DLH;
(g) 16 m2·h
represent 2008 conditions and arrows represent direction and magnitude of change following harvests in 2009. Longer arrows represent
greater difference in the plant community pre- and post-harvest. The solid black ellipsis is associated with the 13-cm DLH. The black dot-
ted ellipsis is associated with the Control. Significant environmental variables can be found in Table 3.

M. A. CAMPIONE ET AL.
Tab
Imprta,
MI. ce
is gr
le 3.
ortant environmental attributes associated with the ordination axes of the NMS used in Figures 3 and 4 at the Ford Forest Cutting Trial, Albe
Environmental variables that were significantly (p = 0.05) correlated to Axis 2 or Axis 3 were included. The relative location in ordination spa
aphed in the inset in Figure 3.
Axis 1 Axis 2 Axis 3
r r2 p value r r2 p value r r2 p value
Average Diameteverstory 0.517 0.0.0010.1960. 0.013 0.48 0.<0.001 r of O267 <03823
Distance to Stream (m) 0.345 0.9 <1 0.70 <1 0.4 0.6 <1
Overstory Basal Area 0.551 0.304 <0.001 0.2860.082 < 0.001 0.58 0.337 <0.001
0.
rsity
0.
11 0.0031 .10.0045 200.00
Saplings Richness 0.06 0.004 0.4510.026 0.001 0.749 308 0.095 <0.001
Saplings Shan no n’s D
Total Seedlings
ive 0.348 0.
0.17
121
0.029
< 0.001
0.032
0.162
0.08 0.
0.026
006
0.04
0.313
0.392 0.
382 0.
154
146
<
<
0.001
0.001
tver, wermanent plots,
composition was not static (FröboOv7).
et been studied extensy es com
erbaceous comShields & Webster, 2007
is in ifting herbacemesfte-
ent s meta-analysis of vscular plan
2 –12 –12 –1
ween 1970 and 1993; howeithin p species
rg &
ivel e, 199 in forThe carou- tsl effect has nomuni-
ties; future monitoring of permanent plots, such as those estab-
lished in this cutting trial, will allow for a more robust assess-
ment of this effect.
Compositional changes amongst the 13-cm diameter-limit
harvest and the control were generally small. The control treat-
ment was generally associated with greater overstory basal area
and larger average diameter of overstory trees compared to the
13-cm diameter-limit harvest, which was associated with
greater richness and diversity in the sapling layer. The uncut
control was dominated mainly by a shade tolerant fern, Dryop-
teris spinulosa, and Acer saccharum seedlings and saplings.
The composition of the 13-cm diameter-limit treatment was
mainly dominated by shade intolerant, exotic species such as
Galeopsis tetrahit, Hieracium aurantiacum, Taraxacum offici-
nale, and Veronica officinalis.
In the 13-cm diameter-limit treatment, Rubus spp. did not
change in frequency but did increase in percent cover. This
increase in percent cover may have contributed to the absence
of the exotic species Anthemis cotula, and Hieracium auran-
tiacum during the 2009 sampling period, all of which are con-
sidered intolerant. Rubus spp. has been shown to delay tree
regeneration and herbaceous establishment in some northern
hardwood forests (Shields and Webster, 2007). Holmes and
Webster (2010) observed in hemlock/hardwood forests a dif-
ference in the herbaceous community in deer access and deer
exclusion plots; plots without fencing (deer access) were gener-
ally dominated by weedy and exotic species. However, interac-
tions between Rubus spp., deer herbivory, and tree regeneration
are often complex and vary regionally (Horsley and Marquis,
1983). Teasing out the effects of management (past and present)
and herbivory on long-term forest dynamics are extremely
complex and often do not have simple answers (Hester et al.,
1996; Gill, 1996).
Following a winter harvest, the diameter-limit treatments had
herbaceous communities that were more similar to the 13-cm
diameter-limit harvest than the control. The directionality and
magnitude of compositional change was consistent across all
diameter-limit treatments. This trend, however, was not ob-
served in the residual basal area treatments. These results did
not support our original hypothesis that harvest intensity, in
terms of the percent overstory basal area removed, would be the
main factor influencing herbaceous composition. However,
changes in light availability may be contributing to changes in
Europe (Paillet et al., 2009). Paillet and colleagues (2009) iden-
tified that species richness was generally higher in managed
forests than unmanaged forests. This trend in greater species
richness was also observed by Scheller and Mladenoff (2002)
in northern hardwood stands located along the border of north-
ern
the hmunity (e.g. ).
Th
agem trend is consish tent withous comuniti a ar man ts in a
Wisconsin and the Upper Peninsula of Michigan. They
observed an increase in species richness in actively managed
stands, both even-aged and uneven-aged, compared to old-
growth stands. Some of the greatest differences in the light
environment, herbaceous abundance, and herbaceous diversity
they observed occurred between uneven-aged and old-growth
stands. The control in our study is not considered old-growth.
However, our control and the even-aged stands in the Scheller
and Mladenoff (2002) study share many similarities and had a
distinctly different herbaceous community than the uneven-
aged stands. The largest environmental difference that Scheller
and Mladenoff (2002) observed in even-aged and old growth
stands was the amount of coarse woody debris. As vegetation
dynamics continue to occur in our control, a more heterogene-
ous environment may occur in the understory which may allow
for the development of a herbaceous layer having more old-
growth qualities.
This shift in herbaceous species composition between dif-
ferent harvest intensities, even-aged and uneven-aged manage-
ment was not observed at the Argonne Experimental Forest in
northern Wisconsin (Kern et al., 2006). The Argonne Experi-
mental Forest includes two diameter-limit treatments, 20-cm
and 30-cm, that were harvested 39 years before vegetation
sampling; three single-tree selection treatments with residual
basal areas of 20.6 m·ha , 17 m·ha , and 13.8 m·ha har-
vested nine years before vegetation sampling; a shelterwood
harvest; and a control. The differences between studies may be
largely due to the difference in exotic species. Within the Ar-
gonne, only three exotic species were observed and all were
considered rare. At the Ford Forest Cutting Trial, on the other
hand, twelve exotic species were observed and some were
rather common (e.g., Galeopsis tetrahit). There are a number of
potential reasons that these two studies have produced seem-
ingly divergent responses. First, propagules may not have been
as common in the surrounding landscape at the time of harvest
or simply failed to invade the site due to barriers to movement
or limited availability. This is a likely casual mechanism since
the Ford Forest Cutting Trial is proximate to a state highway
104

M. A. CAMPIONE ET AL.
and numerous haul roads. Roads are widely recognized vectors
for the spread of exotic species and Buckley et al. (2003) ob-
served that haul roads were one of the primary ways that intro-
duced species enter forested stands in the Upper Peninsula.
Second, the exotic species in question may be transient and
disappear after a suitable recovery period, while remaining
dormant in the seedbank for extended periods (Thompson and
Band, 1997). Short cutting cycles (10 years) at the Ford Forest
Cutting Trial may allow these species to persist and spread
more readily than the longer cutting cycles at the Argonne Ex-
perimental Forest. Third, the pool of potential invaders may be
increasing with time. Consequently, recently applied treatments
may be more susceptible to invasion than older studies were.
Continued treatment of these and other silvicultural trials along
with more consistent monitoring of herbaceous-layer dynamics
would help to shed additional light on the resiliency of forest
plant communities to anthropogenic disturbance and changes in
exotic propagule pressure. Though our results demonstrate the
initial response of the herbaceous community to these treat-
ments, the real strength of this study will be in following the
response of the herbaceous community over time in response to
long-term, consistent silvicultural treatment.
Management Implications
As our understanding of forest ecosystem function expands,
the term sustainable management will continue to include more
complex processes especially with an uncertain climate future.
It is hypothesized that more diverse forest communities may be
more resilient to climate change (reviewed by Hooper et al.,
2005). In a review by Folke et al. (2004), they noted that human
actions can reef-
fects (loss of fu
(e
cts of forest management on
ve
support was provided by the Ecosystem Science Center and the
sources and Environmental Science at
Michigan Technological University, and the McIntire-Stennis
Program.
ion system. 21. Station Paper LS-56. St. Paul, MN: US De-
partment of Agriculture, Forest Service, Lake States Forest Experi-
ment Station.
Berndt. L. W. (1988). ounty area Michigan.
y Management, 175, 509-520.
duce ecosystem resilience through top-down
nctional groups of species), bottom-up effects
nvironmental changes such as climate change and pollutants),
and changes to disturbance regimes. Scheller and Mladenoff
(2002) hypothesize that traditional uneven-aged management
with short cutting cycles may be creating herbaceous communi-
ties dominated by early successional and exotic species. Man-
agers will need to experiment with traditional and non-tradi-
tional techniques to retain and/or enhance native diversity in all
layers of the forest ecosystem. Diameter-limit harvesting is one
traditional technique that may have a greater effect on the
structure of the overstory (Bohn et al., 2011) and in our study
caused a greater shift in the herbaceous layer than traditional
single-tree selection management.
Consistent management in northern hardwood forests is rare.
The Ford Forest Cutting Trial was intended to be a demonstra-
tional forest where scientists and managers could observe the
results of consistent management. Following 50 years of treat-
ment, divergent responses in the herbaceous layer are becoming
apparent along a gradient of harvest intensity. Continual moni-
toring of these treatments will allow future scientists and man-
agers to observe the long-term effe
getation dynamics.
Acknowledgements
The authors are grateful to past Michigan Technological
University professors and staff who have maintained the Ford
Forest Cutting Trial. Wilfred Previant, Jim Schmierer, Jim Ri-
vard, and the FERM team helped with data collection. Financial
School of Forest Re
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illiams, M. (1989). Americans and their forests: A historical geog-
raphy. Cam bridge, UK: Cambridge University Press.

M. A. CAMPIONE ET AL.
Appendix
Table A1.
Complete species list, both scientific and common names, and general chaistics of each species found at the Ford Forest Cutting Trial, Alberta,
MI. USDA Plant Database was used for scientific nomenclature.
ode Name ScieCommon Name
racter
Species Cntific Name
actspp Actaea L. spp Baneberries
adiped Adiantum pe datum L. Northern maidenhair Fer n
alltri Allium tricoccum Aiton Wild leek
Anthemis cotula L.
lis L. illa
.) Schott
Schreb. ex Spreng.) P. Beauv. orthusk
aMuhl. ex Willd. bittercress
ichx.
canadensis (L.) H. Hara ade
a L. m
ld fern
pp.
hempnettle
green
thoides (L.) Sweet eye daisy
m L.
hylla (L.) Pers. f
*
.
L.
x Marsh. honeysuck le
ense Desf. flower
Link n’s seal
eris(L.) Torado
etroot
Michx.) C.B. Clarke
spp. seal
L.
iata E.P. Bi cknell snakeroot
le F.H. Wigg.
Bicknell orse-gentian
s L. edwell
ullein
antcot* Mayweed
aranud Aralia nudic auWild sarsapar
aritri Arisaema triphyl lum (L Jack-in-the-pulpit
braerec Brachyelytrum erectum (Bearded sh
carpen Cardamine pensylvanicPennsylvania
carspp Carex L. spp. Sedge
cautha Caulophyllum thalictroides (L.) MBlue cohos h
cirlut Circaea qu a drisulcata (Maxim.) Franch. & Savigny var.Enchanter’s nightsh
dicspp Dicentra Bernh. spp. Bleeding Heart
diovil Dioscorea villosWild ya
drycar Dryopteris spi nu los a (O.F. Müll.) Watt Spinulose shie
elyhys Elym us hystrix L. Rye grass
eurspp Eurybia (Cass. ) Cass sAster
fraspp Fragaria L. spp. Strawberry
galspp Galium L. spp. Bedstraw
galtet* Galeopsis tetrahit L. Bristlestem
gaupro Gaultheria pr oc um be ns L. Winter
geuspp*Geum L. spp. Avens
helhel Heliopsis helianSmooth ox
hieaur* Hieracium auranti acuOrange hawkweed
ipospp* Ipomoea L. spp. Morning glory
jefdip Jeffersonia dipTwinlea
lapcom Lapsana communis nipplewort
leuvul* Leucanthemum vulgare Lam Oxeye daisy
linbor Linnaea b ore alis Twinflower
loncan Lonicera c anadensis Bartram eAmerican fly
maican Maianthemum canadCanada may
mairac Maianthemum racemosum (L.) False Solomo
matstr Matteuccia strut hioptOstrich fern
menarv Mentha arvensis L. American wild mint
myospp*Myosotis L. spp. Forget-me-nots
oryasp Oryzopsis asperifolia Michx. Rough leaf rice grass
osmchi Osmorhiza chilensis Hook. & Arn. Mountain swe
osmcla Osmorhiza claytoni (Sweet cicely
polspp Polygonatum Mill True Solomon’s
rhacat* Rhamnus cathartica L. Common buckthorn
ribspp Ribes L. spp. Gooseberry
rubspp Rubus L. spp. Raspberry
samspp Sambucus L. spp. Elderberry
sancan Sanguinaria canadensis Bloodroot
santri Sanicula trifolLong-f ruited
staspp Stachys L. spp. Lamb’s ears
taroff* Taraxacum officinaDandelion
triaur Triosteum aurantiacum E.P. Orangefruit h
trispp Trillium L. spp. Trillium
veroff* Veronica officinaliCommon spe
vertha* Verbascum thapsus L. Common m
viospp Viola L. spp. Violet
*Eecies. xotic sp

M. A. CAMPIONE ET AL.
Table A2.
Fr of occurrence oe- and post-harvest by treatment at the Ford Forest Cutting Trial, Aeous species
ararized at the subp0). DLH refers to diameter limit harvest.
Control 21 m2·ha–1
2–1 56 cm DLH 16 m2·ha–1 2–1 41 cm DLH 11 m2·ha–1 2–1 30 cm DLH 13 cm DLH
equencyf herbaceous species prlberta, MI. Herbac
e summlot level (n = 1
(90 ft·ac) (22 in) (70 ft·ac) (16 in) (50 ft·ac) (12 in) (5 in)
Species Co
Name 2008 2009 2008 2009 2008 200920082009
de
2008200920082009 2008 2009 20082009
actspp - - - - - - - - - - 10 - 10 - - -
adiped - - - - - - - - 20 20 - - - - - -
alltri - - -0 - 10 - -
antcot*
b
11 1 1 1
1
1 1
1
1
- - 1- - 1020 60 20
- - - - - - - - - - - - - - 30 -
aranud - - - - 10 10 - 10 - - - - - - - -
aritri - - - - - 10 20 20 30 10 30 10 30 - - -
raerec- - - - - 10 - - - - - 10 - - - -
carpen - - - - - - - - - 20 - - - - - -
carspp 40 - 70 40 80 50 80 60 00 70 90 80 00 00 00 00
cautha - - - - - - 10 10 20 20 50 40 - - - -
cirlut - - - - - - - - - - 10 - - - - -
dicspp - - - - - - - - - 10 - 10 - 10 - -
diovil - - - - - - - - - - - 10 - - - -
drycar 80 90 00 90 80 80 90 90 80 80 80 70 60 40 10 10
elyhys - - - 20 - - - 10 - 40 - 50 - 40 - 90
eurspp - - - - - - - - - - 10 - - - - -
fraspp - - - - - - - - - - - - - - - 20
galspp - - - - - - - - 10 - - - - 20 50 20
galtet* - - 40 60 60 50 80 60 90 100 00100 90 80 00 30
gaupro - - - - - - - - - - - - - - 20 -
geuspp* - - - - - - - - - 20 - 20 - - - -
helhel - - - - - - - - - - - - - - 10 -
hieaur* - - - 10 - - - - - - 10 - - - 50 -
ipospp* - - - - - - 20 - - - - - 20 - 80 -
jefdip - - - - - - 10 - 60 - - - - - - -
lapcom* - - 10 20 20 20 10 20 10 70 90 90 40 40 80 -
leuvul* - - - - - - - - - - - - - - - 20
linbor - - 20 - 10 - - - - - - - - - - -
loncan - 10 - - - 10 - - - - - - - - - -
maican 10 - - - - 10 - - - - - - 10 10 10 40
mairac - 10 - 10 - 10 - - - 30 - - - 10 - -
matstr - - - - - - - - - 10 - - - - - -
menarv - - 10 - - - - - - - - - - - - -
myospp* - - - - - - - - - - - 10 - - - -
oryasp - 30 - 50 - 70 - 60 - 20 - 50 - 70 - -
osmchi - - - - - - - - - 10 - 10 - 20 - -
osmcla - - - - - - - - 30 - - - - - 20 -
polspp - - 10 30 30 10 30 10 50 30 20 30 50 40 - 00
rhacat* - - - - 10 - - - - - - - - - - -
ribspp 30 20 10 10 - 20 - 10 10 10 - 20 50 30 10 -
rubspp - - 90 90 90 90 60 50 60 70 70 80 10 60 100 00
samspp - - - 10 - 30 - 10 - - 10 10 - - - 10
sancan - - - 20 50 50 50 60 80 70 50 80 60 60 20 30
santri - - - - - - - - 10 10 20 - - - - -
staspp - - - - - - - - - - - - - - 10 -
taroff* - - - 10 10 10 - - 20 - 40 30 20 20 70 60
triaur - - - - - - - - 10 10 - - - - - -
trispp 10 10 30 10 20 30 60 40 60 20 80 50 20 40 10 20
veroff* - - - - - 10 - - - - - - - - - 10
vertha* - - - - - - - - - - - - - 20 - -
viospp 10 30 - - 20 20 40 30 40 60 40 40 50 30 70 -
*Exotic ss. pecie
108

M. A. CAMPIONE ET AL.
Tab
Percent of herbaceous species co-ocrrenceetween treatmts in 28 and09 ae Fororestuttingial, Alberta, DLHers i-
ame harves
Control 21 m2·ha–1
(90 ft2·ac–1) 56 cm DLH (22 in) 16 m2·ha–1 (70 ft2·ac–1) 41 cm DLH (16 in) 11 m2·ha–1 (50 ft2·ac–1) 30 cm DLH (12 in) 13 cm DLH (5 in)
le A3. cu ben00 20t thd F C TrMI. refto d
ter limit t.
2008
Control - 33 27 37 25 25 47 32
21 2–1
(90 )
5
(22 in) 27
(70 ft2·ac–1) 37 44 53 - 60 52 61 39
41 cH
(16 in) 25 38 45 60 - 52 52 46
(50 ft2·ac–1) 25 38 45 52 52 - 59 36
30 cH
(12 in) 47 53 53 61 52 59 - 52
(5 in) 32 32 39 39 46 36 52 -
2009 Control 21 m·ha–1
(90 ft2·ac–1) 56 cm DLH (22 in) 16 m·ha–1 (70 ft2·ac–1) 41 cm DLH (16 in) 11 m·ha–1 (50 ft2·ac–1) 30 cm DLH (12 in) 13 cm DLH (5 in)
m·ha
ft2·ac–1 33 - 53 4 438 3 853 32
6 cm DLH53 - 53 45 45 53 39
16 m2·ha–1
m DL
11 m2·ha–1
m DL
13 cm DLH
2 2 2
C10 ontrol - 29 33 28 29 20 29
21 m2–1
(90 )
5
(22 in) 33
(70 ft2·ac–1) 28 63 61 - 54 63 50 41
41 cH
(16 in) 29 50 45 54 - 62 57 30
(50 ft2·ac–1) 20 58 57 63 62 - 59 40
30
(12 in) 29 59 58 50 57 59 - 46
H
(5 in) 10 20 44 41 30 40 46 -
·ha
ft2·ac–1 29 - 57 63 50 58 59 20
6 cm DLH57 - 61 45 57 58 44
16 m2·ha–1
m DL
11 m2·ha–1
cm DLH
13 cm DL