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The Effects of Environmental Variables on Montane Longleaf Pine Ecosystems, Oak
Mountain State Park, Alabama
By:
Kevin Willson
Dr. Scot Duncan and Dr. Malia Fincher, REU Mentors
Samford University
August 2014
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Introduction
Longleaf pine (Pinus palustris) woodlands used to dominate the southeast portion of
the United States. Stretching from eastern Texas to Florida and north to Virginia,
dominant and mixed longleaf pine ecosystems harbored an array of biodiversity and
covered over 37 million hectares of land (Frost 1993, Walker 1984). The longleaf pine
tree grows to the top of the canopy, can live up to 500 years old, and produces a strong,
long-lasting wood that does not decay readily due to the resin found within the tree
(Brockway 1997). The tree is native to regions controlled by fire, which would naturally
sweep through swaths of land every one to three years and clear out the understory of the
woodland (Loudermilk 2011). In the Birmingham area, fires would return every 6-8
years (Bale 2009). Fire creates favorable conditions for longleaf seeds to germinate by
thinning out less fire-resistant plants and trees and exposing seeds to bare mineral soil,
freeing nutrients for immediate consumption, and burning away litter on the forest floor
(Brockway 1997). These mineral soils are exposed by fires burning through most of
these flammable organic soil horizon, burning away the little O and A horizons built up
within the soil (Varner 2005). Other than the ability to resist fire in most life stages,
longleaf does not compete well against other large growing deciduous and pine trees,
especially with anthropogenic changes that have occurred over the past several thousand
years (Landers 1995).
Before European settlers arrived, both nature and Native Americans helped encourage
longleaf pine ecosystems. Lightning would induce fires during the spring and summer,
while Natives introduced additional burnings in the fall and winter to herd larger game
animals for hunting (Frost 2006). European settlers slowly took over the Southeastern
US starting in the 1600s and made large changes to the landscape. People collected the
pine’s resin because it could be converted into turpentine, tar, and pitch, while the wood
was in high demand to construct buildings and lay railroad tracks (Jose 2006). The
tremendous value people placed on longleaf as a resource, coupled with the tree’s
abundance throughout the Southeast, led to immense destruction of the habitats while
supplying an ever growing United States during the 18th, 19th, and 20th centuries. By the
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end of the 1940s, most longleaf stands had been completely logged with little restoration
attempts to regrow the trees (Jose 2006). Starting in the early 1900s, the United States
began implementing a stringent fire suppression regime to try to protect forests, which
unknowingly continued hurting longleaf pine forests. The overharvesting of longleaf
pines paired with intensive fire suppression throughout the South have hurt the species in
its ability to recapture lost land, leading to drastic losses in endemic species needing the
pine and ecosystem to prosper. The lack of fire has allowed the other trees including
loblolly pine and more deciduous trees to outgrow and out-compete the longleaf pine in
most areas throughout the Southeast (Van Lear 2005).
The longleaf pine plays a critical role in maintaining habitats throughout the
Southeast. As a keystone species, the lack of abundant longleaf pine exhibits the
ecosystem’s inability to become properly re-established, indicating the overall loss of
pristine fire climax habitats (Brockway 1997, Landers 1995). The tree’s loss has led to
the demise of a number of plant and animal species, with over 29 plant and animal
species labelled as threatened or endangered because of, or partially due to, the loss of the
longleaf pine ecosystem (Van Lear 2005, Brockway 1997). Montane systems, though
smaller in area, are just as important as its coastal companion in helping restore longleaf
pine. Montane longleaf habitats support different ecosystems than the coastal plains and
could potentially harbor species that are not found elsewhere within the South.
As scientists attempt to restore longleaf pine ecosystems after an era of fire
suppression, researchers, including Van Lear (2005), Varner (2003 & 2005), Brockway
(1997), Lavoie (2010) and Fowler (2007), have studied how this pine species and its
ecosystem have been affected over the past century. The soils of regularly burned
ecosystems feature fluctuating leaf litter and organic soil horizons, which significantly
differs from unburned forest floors that have deeper leaf litter and organic soil horizons
(Lavoie 2010). Montane systems may be unique to other longleaf pine regions because
of differences in slope and soil depth, yet little is known about possible differences
because of the lack of research within mountainous regions of this habitat. Oak
Mountain State Park (OMSP), the research site for the study, is one of the few montane
ecosystems that still contain longleaf pine. Within the field study sites at OMSP, thick
layers of leaf litter are noticeable and have added to soil depth, likely accumulating from
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the lack of consistent fire our research plots have not had over in the recent past. This
soil accumulation affects all ecosystems within the longleaf pine range.
While scientists are collecting large amounts of data on coastal longleaf pine
ecosystems (Drewa 2002, Gitzenstein 2001, Jose 2010, Noel 1998, Outcalt 2010, Peet
1993, etc.), relatively few scientific studies have looked into understanding montane
longleaf ecosystems (Bale 2009, Maceina 2000, Varner 2003). The effects of increased
organic matter in the soil change soil moisture retention, nutrient availability, and soil
bulk density, which may have profound effects on the species richness and abundance
found in the area (Brockway 1997). Though part of the study will look to see how soils
may relate to the number and basal area of adult longleaf pine, we are also interested in
seeing how canopy cover could impact juvenile survival.
Few studies in the past have looked into the effects of canopy cover on longleaf pine
ecosystems. Peacot (2005) found a negative the relationship between the quality of light
coming through canopy and overstory tree stocking, while McGuire (2001) found an
increase in juvenile longleaf pine growth when gaps were created in the canopy from tree
removal. Unfortunately, both of these studies were focused on coastal ecosystems. Our
study will look at the impacts of canopy cover in montane ecosystems in regards to how
light availability affects the growth of juvenile longleaf pine trees.
In this study, we not only attempted to add to the science of montane longleaf
ecosystems, but tried to appreciate how environmental variations within mountainous
areas may change longleaf pine growth. In comparing two different regions of Oak
Mountain State Park, a foothills and a mountain slope zone, we attempted to add
literature of how variations between the smaller foothills and a larger mountain change
the biodiversity and abundance of longleaf pine juveniles and adults. A number of
differences exist between characteristics of foothill and ridge regions in Oak Mountain
State Park including the steepness of the slopes, soil depth, and bedrock variations
between shale (in the foothills) and sandstone (in the ridge). By seeing the change of
habitats between a steeper ridge and rolling foothills, we may better understand the
differences between coastal plain ecosystems, generally flatter than the foothills at
OMSP, and montane ecosystems. We need more data on montane longleaf pine
ecosystems to better understand how the community there interacts with the abiotic
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features of the land and to improve restoration efforts. To learn more about these system,
this study looked into finding patterns that may predict juvenile longleaf pine abundance
and basal area of adult longleaf pine.
Hypotheses: This study looked into two important measures of longleaf pine health,
juvenile longleaf pine abundance and total basal area of adult longleaf pine, and how
several variables may affect and predict these factors. These two measures were studied
because they represent two important factors that help examine both longleaf recruitment
and adult growing capabilities. I hypothesized that variables that decreased soil depth,
increased slope, increased tree species richness, and increased non-longleaf pine basal
area had negative relationships with juvenile longleaf pine frequency and basal area of
adult longleaf pine trees because the less stress and more species of tree filling space and
niches, the fewer number of longleaf will be able to compete within the ecosystem.
I was also interested in determining if a relationship exists between canopy openness
and total number of juvenile longleaf pine in the understory. As a young, small tree,
collecting enough light is one of the most critical components to surviving; the more
light, the more likely a juvenile tree is to survive. Because canopy cover (the opposite of
canopy openness) would affect light availability on the forest floor, I expected canopy
openness and the abundance of juvenile longleaf pine to be mathematically related to one
another and have a positive relationship.
Environmental Variables: Environmental variables measured for this study included
canopy openness, slope steepness and soil depth. Canopy openness, the percentage of
overhead sunlight able to reach the ground, was determined using a hemispherical lens
and camera to take a 180° picture approximately one meter above the ground in the
center of the subplot. The photo was processed using GLA (V. 2) software used to
determine the amount of tree cover within the subplot (Frazer 1999). Slope was recorded
from the lowest to highest points of the perimeter of each subplot and measured with a
clinometer (Suunto PM5/360 PC, Finland). Soil depth was tested at the center of the
subplot as well as at points two meters away from the center of the subplot in the four
cardinal directions. Soil depth was measured by inserting a four foot steel soil probe (3/8
inch diameter made by Forestry Suppliers, Inc in Jackson, MS) into the ground until it
could not go any further or until bedrock was struck and the depth was recorded. The soil
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depth data used for the statistical analyses in this study were the average recordings of the
five measurements in each subplot. Soil depth may affect the total basal area of longleaf
pine in foothill and ridge regions in the park for several reasons. In this region, more
complete organic horizons indicate fewer fires sweeping through the area, which
negatively affects longleaf pines’ ability to compete in the ecosystem. Also, deeper soils
generally have more nutrients and water retention, which allows a larger diversity of trees
and more competition due to less stress in attaining and retaining water and nutrients.
Indicator variables: Indicator variables included total tree species richness, juvenile
longleaf pine abundance, and longleaf pine and non-longleaf pine basal area. Total tree
species per subplot – the species richness - was the unit of measuring biodiversity in this
study. Juvenile longleaf pines shorter than 1.3 meters was measured for basal diameter,
height, and abundance. Only five or six of the ten subplots was measured in each plot (1,
4, 5, 8, 9) and one randomly chosen subplot (2, 3, 6, 7, or 10) if the researcher had
enough time to examine one more subplot. Finally, we looked at tree biodiversity and
basal area measured in these plots by Dr. Scot Duncan ten years ago to try finding
possible variables that affect tree biodiversity – measured by recording total number of
tree species in each subplot – and total basal area of longleaf pines, which was calculated
by taking the area of the tree at breast height. Though some comparisons used in the
research were taken from data collected ten years apart, the slope and soil depth data that
we collected in 2014 would not likely change much from 2004.
The statistical analyses were performed using SPSS (Version 19, IBM). The Mann-
Whitney U non-parametric test provided the statistical evidence of differences between
the ridge and foothills as well as differences between subplots with and without longleaf
pine adults/juveniles. Multiple linear regressions were used to determine how predictive
the independent measured variables were on the dependent variables (juvenile longleaf
pine frequency and adult longleaf pine basal area per plot). All of the data was tested
using multiple linear regressions and then subplots with no longleaf pine were taken out
of the analysis for the purposes of seeing: when longleaf pine did grow, what were factors
that may influence growth. If the p value was under 0.200, the variable was kept to look
in the next multiple linear regression test to see if it became significant.
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Methods
Study Site: Oak Mountain State Park is located in Pelham, Alabama and contains
9,940 acres of land (Alabama State Parks Website). Numerous ecosystems are found in
the park, from the ridgeline woodland at the top of Double Oak Mountain to the foothill
forests and stream habitats that are scattered around the mountain. Double Oak Mountain
is a twin ridged, ridge and valley system that is runs in a Northeast to Southwest
direction. The mountain influences many aspects of the natural ecology in the park from
water runoff to plant species diversity found in various areas within OMSP. The park
features two different topographic regions due to the mountain, including the ridge (the
mountain face) and the foothills, the rolling hills around the mountain. The ridge is
higher in altitude, topping out at 1,260 feet above sea level and is made mostly of shale in
the mountain’s valley and sandstone in the twin peaks. The foothills fluctuate in
topography greatly, having numerous high points as compared to the ridge which only
has several ridgeline high points and long slopes. The foothills are also mostly made of
shale with traces of sandstone. In the past, most of this land likely contained a large
amount of longleaf pine judging by the number of longleaf pine stumps still found on the
ridge and in the foothills, though other trees have overtaken many parts of the park.
Average rainfall in the area is approximately 135 centimeters of rain per year (Maceina
2000) and average high temperatures range from 54 °F in January to 91 °F in July
(Birmingham). Frost and freezing temperatures occur between one to three times per
year (Maceina 2000).
Study Design: The research plots used as the study sites at Oak Mountain State Park
were created approximately 11 years ago by Dr. Scot Duncan. Ten foothill plots were
randomly placed on the top of forested hills within the foothills region of the park. Ten
ridge plots were selected at random on the southeast slopes of Double Oak Mountain.
These plots were 50x20 meters and divided into ten 10x10 meter subplots. Each subplot
was individually measured for the study’s variables.
Results
Juvenile longleaf pine were not found very frequently throughout the subplots as
compared to adult longleaf pine, only found in 34% of the 108 subplots sampled versus
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adult longleaf found in 64.5% of 200 subplots. Within the foothill region, four plots had
juvenile longleaf for a grand total of five subplots (9.09% of sampled subplots). In the
ridge, all 10 plots had longleaf pine juveniles for a total of 32 (60.3%) of the 53 subplots
sampled. The ridge subplots that had juvenile longleaf averaged 12.2 juveniles per
subplot, whereas the foothills plots averaged 2.8 per subplot.
Longleaf pine tree basal area found within the plots significantly varied between and
within the foothills and ridge. Within the foothills plots, there was an average of 31
longleaf pine trees per plot, making up approximately 18% of the numbers of trees
recorded, while the ridge plots contained an average of 14.9 longleaf pine per plot (19%
of the number of trees). Total longleaf basal area in subplots had less variability than in
the foothills, while the ridge longleaf pines averaged a larger basal area overall (Table 1).
The total basal area of other trees in the foothill subplots totaled about 15.64 m2, while
ridge plots totaled about 7.93 m2. Tree species totals found that foothill subplots
harbored roughly 1.5 more species overall per subplot than as compared to the ridge
subplots (Table 1).
Canopy openness was relatively similar between the foothill and ridge plots, though
the foothill plots kept a relative uniform average while the ridge plots saw more variation
(Table 1). Slope had significant differences between the two regions and was greater on
ridge subplots. The soil depth average was almost two times deeper in the foothill plots
than as compared to the ridge plots. Significant portions of the subplots in the ridge
lacked canopy cover due to very shallow or absent soils, which never occurred in the
foothills.
Four independent variables (slope, soil depth, species richness, and non-longleaf pine
basal area) were used in a comparison of subplots with and without adult longleaf pine in
ridge plots. In these tests, the only significant variable differing between the two sets of
plots was non-longleaf pine total basal area, which doubled in subplots without longleaf
pine (Table 4). In the foothill plots, the only two significantly different variables between
subplots with and without longleaf pine were the species richness and non-longleaf tree
total basal area variables.
In a similar comparison between subplots with and without juvenile longleaf pine,
four different variables (canopy openness, slope, soil depth, and tree species richness)
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were significantly different between subplots. In the foothills, slope was much shallower
and soil depth was much deeper than in the ridge, as seen in Table 3. The ridge subplot
comparison found that canopy openness and soil depth were the two variables
significantly different in subplots with and without the juveniles (Table 4).
Multiple Regression Models:
In the first regression model, juvenile longleaf pine frequency was the dependent
variable and canopy openness, soil depth, tree species richness, slope, and non-longleaf
pine basal area. The data was separated by foothills and ridge. Running a multiple linear
regression test for foothill plots found that the three variables tested (slope, soil depth,
and species richness, excluding non-longleaf pine basal area and canopy openness due to
lack of statistical strength) were all significant in predicting juvenile frequency within the
five subplots the juveniles were found in (Table 2). Analysis of the ridge plots found that
soil depth, non-longleaf pine basal area, and slope did not play a significant role in
predicting juvenile frequency, but canopy openness and species richness did positively
correlate to the is variable significantly.
The next question looked for a connection between the dependent variables of
biodiversity and longleaf pine basal area and the independent variables of soil depth,
slope, tree species richness, and non-longleaf pine basal area. Subplots that did not
contain longleaf pine were removed and the regression was run again with the same
dependent and independent variables. In the ridge plots, the significant variables in a
multiple linear regression included soil depth, slope, and “other” tree total basal area. In
a multiple linear regression only including these three variables, the test found a strong
correlation to predicting longleaf pine basal area (R2 = 0.363, p < 0.001). Individually,
slope was negatively predicted basal area, the soil depth positively predicted basal area,
and the non-longleaf pine total basal area negatively predicted longleaf pine basal area
(Table 2).
Discussion
Overall, we found that the biggest factors we tested affecting longleaf pine growth in
Oak Mountain State Park were stress and competition. Stress was best measured through
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two environmental stressors, slope and soil depth, competition was measured with a
comparison of total basal areas of trees, and stress and competition were measured
together by tree species richness. Our findings suggest that the longleaf pine have had to
deal with significantly different environmental conditions within the two subsets of a
montane ecosystem. The ridge is a more difficult and stressful environment for plants to
grow as compared to the foothills. Also, succession due to fire suppression is more
advanced in the foothills than in the ridge because invading tree species have an easier
time getting a foothold there due to less stress.
The ridge and foothill regions of Oak Mountain State Park appear to be very different
environmentally, with significantly different slopes, soil depths, tree species richness and
non-longleaf pine basal area. All of these factors seemed to have played a role in
affecting the growth of juvenile and adult longleaf pine.
Juvenile longleaf pine, if present, would indicate that the longleaf pines in the forest
are mature enough to reproduce and the there are enough beneficial factors to allow the
seeds to germinate into the juvenile stage. The differences in the variables measured
must have affected regeneration of longleaf pine, seeing as regeneration nearly halted in
the foothills, yet recruitment of seedlings continued on the ridge. Due to the lack of
juvenile longleaf pine in the foothill subplots, the results given have to be taken as
provincial findings due to small sample sizes, though this lack of juveniles in the foothills
also provides evidence that juvenile longleaf pine are more likely to survive in more
stressful environments due to lack of competition for space and resources. This disparity
exists despite the fact that adult longleaf basal area was higher in the foothills.
Adult longleaf pine basal area was used as an indirect measure of total LLP space and
indicates the total space and resources taken up by this one species in a subplot. There
was significantly more longleaf pine basal area in the foothill plots, meaning there was
more successful growth of longleaf pine overall in the foothill plots than in the ridge
plots.
Environmental variables seem to directly impact growth of the longleaf pine
community both on individual and collective levels. Slope was steeper in the ridge than
in the foothills. Differences in slope are due to the geology of the mountain and foothills,
where ridge plots are above a thick layer of sandstone, which is not as easily eroded as
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the shale in the rest of the park. Sandstone prevents groundwater penetration for long
term storage, hurting trees during drier periods, while steeper slopes quicken water runoff
and groundwater flow. Juvenile longleaf pine were not significantly affected by the
indirect effects of slope in the ridge, nor were there any differences in ridge subplots with
and without juvenile longleaf pine. In the foothills, slope was negatively related to
juvenile longleaf pine frequency, with significantly less slope when juvenile longleaf was
present in the foothills. For adults, steeper slopes on the ridge had a negative relationship
for longleaf pine basal area, though no pattern was found in the foothill plots between the
two variables. Subplots with adult longleaf pine and subplots without longleaf pine did
not differ for slope in either the foothills or on the ridge. Combined, these two results
indicate shallower slopes generally increased longleaf pine growth, but was not the
variable that allowed or prevented the establishment of the adult tree. Overall, when
slope causes enough stress, it affects the tree to continue to grow, with tree growth
improving on shallower slope. The data show that steeper slopes may hinder all species
of tree growth, but could benefit juvenile longleaf pine due to less competition. The data
also indicates that if slope was steep enough, then it could indirectly affect tree growth
and therefore may affect the longleaf pine community.
Soil depth was deeper in the foothills than in the ridge and may be a pivotal variable in
affecting longleaf pine recruitment and growth. In foothills, juvenile longleaf pine numbers
had a significant negative relationship to soil depth, but no such relationship existed in the
ridge. In the foothills, soil depth was almost a perfect predictor of juvenile longleaf pine
frequency and count, showing strong trends that stress in the habitat may be beneficial for
longleaf pine to become established. Adult longleaf pine basal area in the ridge was
marginally non-significant, but positively related to soil depth, with deeper soils indicating
greater basal area, while there was no significant trend found in the foothills. There was also
no significant difference in subplots where adult longleaf pine was present or absent in either
the foothills or the ridge. The lack of change between subplots with and without adult
longleaf pine indicates that soil depth was not the variable that allowed or prevented the
establishment of the adult tree. When compiling all of this data, soil depth provides evidence
that it causes stress that affects longleaf pines’ continued growth when shallow enough and
that the deeper the soil, the easier for the tree to grow. Like with slope, if soil depth was not
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extreme enough, then it would not play a direct factor in the longleaf pine community.
Finally, longleaf pine generally grows better in deeper soils, but the juvenile longleaf pine
have more success in shallower soils due to less competition. The shallower soil may be a
limiting factor in survival for other plants, but not for juvenile longleaf pine; this opens up
space and resources for more juvenile longleaf pine to germinate and grow.
Non-longleaf pine basal area, being greater in the foothills than the in the ridge, seemed
to have a negative impact on the ability for adult longleaf pine to grow. Non-longleaf pine
basal area was used as a way to measure the amount of space non-longleaf pine trees took in
the two regions of the park, giving an idea of the total area of trees are found in each subplot.
There was significantly more tree area in the foothills than as compared to the ridge,
supporting that the notion that higher amounts of stress lead to lower amounts of total tree
growth. Overall, the only significant regression result for both juvenile and adult growth was
for adult longleaf pine basal area in the ridge, which has a negative relationship. There was
also a significant drop in non-longleaf pine basal area when longleaf pine were in the area.
This supports the idea that other trees generally compete against and are bad for the LLP
community. With more stress, there will be both lower non-longleaf pine basal area and
higher longleaf pine take up space per subplot. Combined with tree species richness results,
the data may provide evidence that that non-longleaf pine trees and the longleaf pine
community have an antagonistic relationship.
Tree species richness was also greater in the foothills than in the ridge. Tree species
richness was used as an indicator of the state of the ecosystem, with increased richness
meaning greater competition for longleaf pine trees. This variable also pointed to possible
stress variations between ridge and foothills. In the ridge, tree species richness was
negatively related to longleaf pine basal area, meaning increased diversity related to the
declining health of longleaf pine community, while decrease in diversity would be beneficial
to longleaf pine trees. Tree species richness was greater in foothill subplots without adult
longleaf pine, supporting the idea that as species richness increases, the more competition
and the worse it is for the longleaf pine community. The ridge plots were not significantly
affected by species richness, which due to the lesser amounts competition from increased
stress. For juvenile longleaf pine, tree species richness was positively related in the ridge,
but negatively related in the foothills. The mixed results support the idea that lower levels of
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stress in a high stress environment create higher chances of germination and growth into the
juvenile stages. In the foothills, more stress is needed to weed out competition for juvenile
longleaf pine to grow, so more stress means less competition. The findings suggest that
longleaf pine compete with other tree species for resources: the more resources the longleaf
pine obtain, the more area they are able to take. Juvenile longleaf also have to compete with
other species when there is little stress, but when stress is elevated to affect the amount of
species, juvenile longleaf pine only need to worry about stressors like soil depth and slope.
We were very surprised to find that canopy openness lacked any significant trends
with any of the tests performed. Young longleaf trees need many hours of direct sunlight
to survive and grow each day. Canopy openness is directly related to light availability at
the forest floor. When more light allowed to reach the forest floor, undergrowth receive
more energy to photosynthesize and produce sugars to store, making it an important
factor in small plant growth. There was one marginally non-significant result when
comparing canopy openness with juvenile longleaf pine in the ridge, with a positive
relationship between the two. The number of insignificant results provide evidence that
canopy openness did not play a major role in longleaf pine recruitment or growth. This
lack of difference may exist because both areas may be at or close to full closure, which
is especially true in the foothills with an increased invasion of broadleaf trees.
Conclusion:
Stress is helpful for LLP as steeper slopes and shallower soils introduce stress that has
been waning with fire suppression. The lack of stress has allowed increased competition
against the longleaf pine, with elevated tree species richness and growth of non-longleaf pine
negatively and unevenly impacting this community the foothills region of the park. These
have seemed to increased stress in the longleaf pine community to have a greater negative
impact than slope and soil depth do. Both of these could be key factors in successful
germination, with soil depth being significantly lower and slope being significantly greater
when comparing plots with and without juvenile LLP. Using these initial findings, we can
start looking into how these variables predict and/or affect the ability for this tree to establish
itself from the juvenile stage into an adult and how it continues to grow throughout its
lifetime.
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In future research, other factors that should be looked into include temperature, slope
aspect, and the average distance away from the parent tree. All of these could also be
factors that cause changes in the abundance of juvenile longleaf pine and size of adult
LLP in each of the subplots. For example, ridge slopes face southeast, meaning the
summer sun is more intense on the slope than others facing different directions, drying
the soils more quickly than on the north facing slopes. How does this change with
different slope aspects? Chance also may play a large factor of deciding whether a
juvenile or adult longleaf pine will grow from a seed into a juveniles or juveniles into an
adult, respectively.
Using these initial findings, we can start looking into how these variables predict
and/or affect the ability for this tree to germinate and grow from seeds into the juvenile
stage and into an adult. There is generally poor recruitment of longleaf pine in the park
overall, so how can that be improved upon by the trees? With this poor recruitment
means a potential drop in adult longleaf pine in the future. With this initial research, we
will need to look for more a detailed understanding of what other environmental factors
play a role in the growth of longleaf pine to help them maintain and re-extend their
community in the future.
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Tables and Graphs
Table 1. Resultsof a Mann-WhitneyUtestseeingif there were differencesbetweenthe ridge and
foothill regionsin OakMountainState Parkin Pelham, AL. A p value of lessthan.05 indicatesa
significantdifference.
Foothill Ridge
Variable Mean (SD) Range Mean (SD) Range P value
Canopy Openness (%) 12.7 (1.7) 7.9 – 17.5 14.5 (5.5) 9.5 – 36.7 0.176
Slope (⁰) 12.0 (5.8) 1.0 – 28.0 21.0 (6.0) 5.0 – 41.0 <0.001
Avg Soil Depth (cm) 34.2 (13.9) 4.9– 65.2 18.3 (13.9) 0.0 – 53.0 <0.001
Juvenile LLP Count 2.8 (1.8) 1.0 – 5.0 12.2 (16.3) 1.0 – 53.0 <0.001
LLP BasalArea (m2
) 0.18 (.15) 0.0024 - 0.84 0.11 (0.09) 0.0005 -0.47 0.004
Tree Species Richness 5.6 (2.0) 2.0 – 11.0 4.1 (1.5) 1.0 - 9.0 <0.001
Non-LLP BasalArea (m2
) 0.13 (.09) 0.016 - .51 0.065 (.06) 0.0009 - 0.33 <0.001
Table 2. Resultsof multiple linearregressionsof the numberof juvenile longleaf pine oradultlongleaf
pine basal area(dependentvariables) againstCanopyOpenness(and/orSpeciesrichness),Slope,basal
area of Non-longleaf trees,andSoil Depth(independentvariables) intwodifferent topographicregions
of OakMountainState Parkin Pelham,AL.Pvalueslessthan.05 representsignificantR2
values.
Region Model’sDependent
variable
R2 P Independent
variables
Beta P
Foothills JuvenileFrequency 1.000 0.011 Slope -0.442 0.020
Soil Depth -0.953 0.006
SpeciesRichness -0.232 0.038
AdultBasal Area 0.204 0.030 CanopyOpenness 0.224 0.124
Slope 0.144 0.281
Soil Depth -0.105 0.438
SpeciesRichness -0.214 0.139
Non-LLPBasal Area -0.132 0.393
Ridge JuvenileFrequency 0.355 0.001 CanopyOpenness 0.313 0.067
Slope -0.274 0.145
Soil Depth 0.065 0.845
SpeciesRichness 0.537 <0.001
Non-LLPBasal Area -0.105 0.431
AdultBasal Area 0.387 <0.001 CanopyOpenness -0.176 0.115
Slope -0.386 0.001
Soil Depth 0.208 0.068
SpeciesRichness -0.024 0.824
Non-LLPBasal Area -0.280 0.014
Table 3. Comparisonof subplotswithandwithoutjuvenile longleaf pineinOakMountainState Parkin
Pelham,AL.Pvalueslessthan.05 representasignificant difference betweenthe subplotsusingaMann-
WhitneyU test.
With Juvenile LLP Without Juvenile LLP
Region Variables Mean (SD) Range Mean (SD) Range P Value
Foothills CanopyOpenness(%) 12.5 (1.7) 10.2 – 14.0 13.9 (5.0) 9.6 – 32.8 0.447
16
Slope (⁰) 12.6 (5.8) 7.0 – 28.0 19.9 (5.7) 7.0 – 32.0 0.002
AvgSoil Depth(cm) 32.52 (14.0) 15.7 – 54.2 23.9 (13.3) 0.0 – 48.1 0.002
SpeciesRichness 4.8 (2.0) 2.0 – 7.0 5.8 (1.9) 3.0 – 11.0 0.437
Ridge CanopyOpenness(%) 14.4 (5.5) 9.8 – 36.7 13.9 (5.0) 9.6 – 32.8 0.466
Slope (⁰) 21.5 (6.0) 9.0 - 38.0 19.9 (5.7) 7.0 – 31.5 0.357
AvgSoil Depth(cm) 16.8 (13.7) 0.0 – 48.4 23.9 (13.3) 0.0 – 48.1 0.103
SpeciesRichness 3.7 (1.3) 2.0 – 6.0 4.3 (1.4) 1.0 – 9.0 0.155
Table 4. Comparisonof subplotswith andwithoutadultlongleaf pineinOakMountainState Parkin
Pelham,AL.Pvalueslessthan.05 representasignificantdifference betweenthe subplotsusingaMann-
WhitneyU test.
With adult LLP Without adult LLP
Region Variables Mean (SD) Range Mean (SD) Range P Value
Foothills Slope (⁰) 12.5 (5.7) 2.0 – 28.0 11.4 (5.8) 1.0 – 26.0 0.423
AvgSoil Depth(cm) 34.1 (13.8) 4.9 – 64.4 34.2 (14.4) 12.3 – 65.2 0.908
SpeciesRichness 5.6 (1.9) 2.0 – 11.0 5.73 (1.9) 3.0 – 11.0 <0.001
Non-LLPBasal Area(m2
) 0.13 (0.09) 0.016 – 0.51 .20 (0.09) 0.070 - 0.44 <0.001
Ridge Slope (⁰) 21.0 (6.0) 5.0 – 41.0 21.1 (5.8) 8.0 – 32.0 0.886
AvgSoil Depth(cm) 17.8 (13.8) 0.0 – 48.4 19.5 (13.3) 0.0 – 48.1 0.564
SpeciesRichness 4.14 (1.4) 1.0 – 9.0 3.1 (1.26) 2.0 – 5.0 0.966
Non-LLPBasal Area(m2
) 0.065 (0.063) 0.0 – 0.22 0.11 (0.064) 0.017 - 0.33 <0.001
17
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Longleaf Pine Final Paper

  • 1. 1 The Effects of Environmental Variables on Montane Longleaf Pine Ecosystems, Oak Mountain State Park, Alabama By: Kevin Willson Dr. Scot Duncan and Dr. Malia Fincher, REU Mentors Samford University August 2014
  • 2. 2 Introduction Longleaf pine (Pinus palustris) woodlands used to dominate the southeast portion of the United States. Stretching from eastern Texas to Florida and north to Virginia, dominant and mixed longleaf pine ecosystems harbored an array of biodiversity and covered over 37 million hectares of land (Frost 1993, Walker 1984). The longleaf pine tree grows to the top of the canopy, can live up to 500 years old, and produces a strong, long-lasting wood that does not decay readily due to the resin found within the tree (Brockway 1997). The tree is native to regions controlled by fire, which would naturally sweep through swaths of land every one to three years and clear out the understory of the woodland (Loudermilk 2011). In the Birmingham area, fires would return every 6-8 years (Bale 2009). Fire creates favorable conditions for longleaf seeds to germinate by thinning out less fire-resistant plants and trees and exposing seeds to bare mineral soil, freeing nutrients for immediate consumption, and burning away litter on the forest floor (Brockway 1997). These mineral soils are exposed by fires burning through most of these flammable organic soil horizon, burning away the little O and A horizons built up within the soil (Varner 2005). Other than the ability to resist fire in most life stages, longleaf does not compete well against other large growing deciduous and pine trees, especially with anthropogenic changes that have occurred over the past several thousand years (Landers 1995). Before European settlers arrived, both nature and Native Americans helped encourage longleaf pine ecosystems. Lightning would induce fires during the spring and summer, while Natives introduced additional burnings in the fall and winter to herd larger game animals for hunting (Frost 2006). European settlers slowly took over the Southeastern US starting in the 1600s and made large changes to the landscape. People collected the pine’s resin because it could be converted into turpentine, tar, and pitch, while the wood was in high demand to construct buildings and lay railroad tracks (Jose 2006). The tremendous value people placed on longleaf as a resource, coupled with the tree’s abundance throughout the Southeast, led to immense destruction of the habitats while supplying an ever growing United States during the 18th, 19th, and 20th centuries. By the
  • 3. 3 end of the 1940s, most longleaf stands had been completely logged with little restoration attempts to regrow the trees (Jose 2006). Starting in the early 1900s, the United States began implementing a stringent fire suppression regime to try to protect forests, which unknowingly continued hurting longleaf pine forests. The overharvesting of longleaf pines paired with intensive fire suppression throughout the South have hurt the species in its ability to recapture lost land, leading to drastic losses in endemic species needing the pine and ecosystem to prosper. The lack of fire has allowed the other trees including loblolly pine and more deciduous trees to outgrow and out-compete the longleaf pine in most areas throughout the Southeast (Van Lear 2005). The longleaf pine plays a critical role in maintaining habitats throughout the Southeast. As a keystone species, the lack of abundant longleaf pine exhibits the ecosystem’s inability to become properly re-established, indicating the overall loss of pristine fire climax habitats (Brockway 1997, Landers 1995). The tree’s loss has led to the demise of a number of plant and animal species, with over 29 plant and animal species labelled as threatened or endangered because of, or partially due to, the loss of the longleaf pine ecosystem (Van Lear 2005, Brockway 1997). Montane systems, though smaller in area, are just as important as its coastal companion in helping restore longleaf pine. Montane longleaf habitats support different ecosystems than the coastal plains and could potentially harbor species that are not found elsewhere within the South. As scientists attempt to restore longleaf pine ecosystems after an era of fire suppression, researchers, including Van Lear (2005), Varner (2003 & 2005), Brockway (1997), Lavoie (2010) and Fowler (2007), have studied how this pine species and its ecosystem have been affected over the past century. The soils of regularly burned ecosystems feature fluctuating leaf litter and organic soil horizons, which significantly differs from unburned forest floors that have deeper leaf litter and organic soil horizons (Lavoie 2010). Montane systems may be unique to other longleaf pine regions because of differences in slope and soil depth, yet little is known about possible differences because of the lack of research within mountainous regions of this habitat. Oak Mountain State Park (OMSP), the research site for the study, is one of the few montane ecosystems that still contain longleaf pine. Within the field study sites at OMSP, thick layers of leaf litter are noticeable and have added to soil depth, likely accumulating from
  • 4. 4 the lack of consistent fire our research plots have not had over in the recent past. This soil accumulation affects all ecosystems within the longleaf pine range. While scientists are collecting large amounts of data on coastal longleaf pine ecosystems (Drewa 2002, Gitzenstein 2001, Jose 2010, Noel 1998, Outcalt 2010, Peet 1993, etc.), relatively few scientific studies have looked into understanding montane longleaf ecosystems (Bale 2009, Maceina 2000, Varner 2003). The effects of increased organic matter in the soil change soil moisture retention, nutrient availability, and soil bulk density, which may have profound effects on the species richness and abundance found in the area (Brockway 1997). Though part of the study will look to see how soils may relate to the number and basal area of adult longleaf pine, we are also interested in seeing how canopy cover could impact juvenile survival. Few studies in the past have looked into the effects of canopy cover on longleaf pine ecosystems. Peacot (2005) found a negative the relationship between the quality of light coming through canopy and overstory tree stocking, while McGuire (2001) found an increase in juvenile longleaf pine growth when gaps were created in the canopy from tree removal. Unfortunately, both of these studies were focused on coastal ecosystems. Our study will look at the impacts of canopy cover in montane ecosystems in regards to how light availability affects the growth of juvenile longleaf pine trees. In this study, we not only attempted to add to the science of montane longleaf ecosystems, but tried to appreciate how environmental variations within mountainous areas may change longleaf pine growth. In comparing two different regions of Oak Mountain State Park, a foothills and a mountain slope zone, we attempted to add literature of how variations between the smaller foothills and a larger mountain change the biodiversity and abundance of longleaf pine juveniles and adults. A number of differences exist between characteristics of foothill and ridge regions in Oak Mountain State Park including the steepness of the slopes, soil depth, and bedrock variations between shale (in the foothills) and sandstone (in the ridge). By seeing the change of habitats between a steeper ridge and rolling foothills, we may better understand the differences between coastal plain ecosystems, generally flatter than the foothills at OMSP, and montane ecosystems. We need more data on montane longleaf pine ecosystems to better understand how the community there interacts with the abiotic
  • 5. 5 features of the land and to improve restoration efforts. To learn more about these system, this study looked into finding patterns that may predict juvenile longleaf pine abundance and basal area of adult longleaf pine. Hypotheses: This study looked into two important measures of longleaf pine health, juvenile longleaf pine abundance and total basal area of adult longleaf pine, and how several variables may affect and predict these factors. These two measures were studied because they represent two important factors that help examine both longleaf recruitment and adult growing capabilities. I hypothesized that variables that decreased soil depth, increased slope, increased tree species richness, and increased non-longleaf pine basal area had negative relationships with juvenile longleaf pine frequency and basal area of adult longleaf pine trees because the less stress and more species of tree filling space and niches, the fewer number of longleaf will be able to compete within the ecosystem. I was also interested in determining if a relationship exists between canopy openness and total number of juvenile longleaf pine in the understory. As a young, small tree, collecting enough light is one of the most critical components to surviving; the more light, the more likely a juvenile tree is to survive. Because canopy cover (the opposite of canopy openness) would affect light availability on the forest floor, I expected canopy openness and the abundance of juvenile longleaf pine to be mathematically related to one another and have a positive relationship. Environmental Variables: Environmental variables measured for this study included canopy openness, slope steepness and soil depth. Canopy openness, the percentage of overhead sunlight able to reach the ground, was determined using a hemispherical lens and camera to take a 180° picture approximately one meter above the ground in the center of the subplot. The photo was processed using GLA (V. 2) software used to determine the amount of tree cover within the subplot (Frazer 1999). Slope was recorded from the lowest to highest points of the perimeter of each subplot and measured with a clinometer (Suunto PM5/360 PC, Finland). Soil depth was tested at the center of the subplot as well as at points two meters away from the center of the subplot in the four cardinal directions. Soil depth was measured by inserting a four foot steel soil probe (3/8 inch diameter made by Forestry Suppliers, Inc in Jackson, MS) into the ground until it could not go any further or until bedrock was struck and the depth was recorded. The soil
  • 6. 6 depth data used for the statistical analyses in this study were the average recordings of the five measurements in each subplot. Soil depth may affect the total basal area of longleaf pine in foothill and ridge regions in the park for several reasons. In this region, more complete organic horizons indicate fewer fires sweeping through the area, which negatively affects longleaf pines’ ability to compete in the ecosystem. Also, deeper soils generally have more nutrients and water retention, which allows a larger diversity of trees and more competition due to less stress in attaining and retaining water and nutrients. Indicator variables: Indicator variables included total tree species richness, juvenile longleaf pine abundance, and longleaf pine and non-longleaf pine basal area. Total tree species per subplot – the species richness - was the unit of measuring biodiversity in this study. Juvenile longleaf pines shorter than 1.3 meters was measured for basal diameter, height, and abundance. Only five or six of the ten subplots was measured in each plot (1, 4, 5, 8, 9) and one randomly chosen subplot (2, 3, 6, 7, or 10) if the researcher had enough time to examine one more subplot. Finally, we looked at tree biodiversity and basal area measured in these plots by Dr. Scot Duncan ten years ago to try finding possible variables that affect tree biodiversity – measured by recording total number of tree species in each subplot – and total basal area of longleaf pines, which was calculated by taking the area of the tree at breast height. Though some comparisons used in the research were taken from data collected ten years apart, the slope and soil depth data that we collected in 2014 would not likely change much from 2004. The statistical analyses were performed using SPSS (Version 19, IBM). The Mann- Whitney U non-parametric test provided the statistical evidence of differences between the ridge and foothills as well as differences between subplots with and without longleaf pine adults/juveniles. Multiple linear regressions were used to determine how predictive the independent measured variables were on the dependent variables (juvenile longleaf pine frequency and adult longleaf pine basal area per plot). All of the data was tested using multiple linear regressions and then subplots with no longleaf pine were taken out of the analysis for the purposes of seeing: when longleaf pine did grow, what were factors that may influence growth. If the p value was under 0.200, the variable was kept to look in the next multiple linear regression test to see if it became significant.
  • 7. 7 Methods Study Site: Oak Mountain State Park is located in Pelham, Alabama and contains 9,940 acres of land (Alabama State Parks Website). Numerous ecosystems are found in the park, from the ridgeline woodland at the top of Double Oak Mountain to the foothill forests and stream habitats that are scattered around the mountain. Double Oak Mountain is a twin ridged, ridge and valley system that is runs in a Northeast to Southwest direction. The mountain influences many aspects of the natural ecology in the park from water runoff to plant species diversity found in various areas within OMSP. The park features two different topographic regions due to the mountain, including the ridge (the mountain face) and the foothills, the rolling hills around the mountain. The ridge is higher in altitude, topping out at 1,260 feet above sea level and is made mostly of shale in the mountain’s valley and sandstone in the twin peaks. The foothills fluctuate in topography greatly, having numerous high points as compared to the ridge which only has several ridgeline high points and long slopes. The foothills are also mostly made of shale with traces of sandstone. In the past, most of this land likely contained a large amount of longleaf pine judging by the number of longleaf pine stumps still found on the ridge and in the foothills, though other trees have overtaken many parts of the park. Average rainfall in the area is approximately 135 centimeters of rain per year (Maceina 2000) and average high temperatures range from 54 °F in January to 91 °F in July (Birmingham). Frost and freezing temperatures occur between one to three times per year (Maceina 2000). Study Design: The research plots used as the study sites at Oak Mountain State Park were created approximately 11 years ago by Dr. Scot Duncan. Ten foothill plots were randomly placed on the top of forested hills within the foothills region of the park. Ten ridge plots were selected at random on the southeast slopes of Double Oak Mountain. These plots were 50x20 meters and divided into ten 10x10 meter subplots. Each subplot was individually measured for the study’s variables. Results Juvenile longleaf pine were not found very frequently throughout the subplots as compared to adult longleaf pine, only found in 34% of the 108 subplots sampled versus
  • 8. 8 adult longleaf found in 64.5% of 200 subplots. Within the foothill region, four plots had juvenile longleaf for a grand total of five subplots (9.09% of sampled subplots). In the ridge, all 10 plots had longleaf pine juveniles for a total of 32 (60.3%) of the 53 subplots sampled. The ridge subplots that had juvenile longleaf averaged 12.2 juveniles per subplot, whereas the foothills plots averaged 2.8 per subplot. Longleaf pine tree basal area found within the plots significantly varied between and within the foothills and ridge. Within the foothills plots, there was an average of 31 longleaf pine trees per plot, making up approximately 18% of the numbers of trees recorded, while the ridge plots contained an average of 14.9 longleaf pine per plot (19% of the number of trees). Total longleaf basal area in subplots had less variability than in the foothills, while the ridge longleaf pines averaged a larger basal area overall (Table 1). The total basal area of other trees in the foothill subplots totaled about 15.64 m2, while ridge plots totaled about 7.93 m2. Tree species totals found that foothill subplots harbored roughly 1.5 more species overall per subplot than as compared to the ridge subplots (Table 1). Canopy openness was relatively similar between the foothill and ridge plots, though the foothill plots kept a relative uniform average while the ridge plots saw more variation (Table 1). Slope had significant differences between the two regions and was greater on ridge subplots. The soil depth average was almost two times deeper in the foothill plots than as compared to the ridge plots. Significant portions of the subplots in the ridge lacked canopy cover due to very shallow or absent soils, which never occurred in the foothills. Four independent variables (slope, soil depth, species richness, and non-longleaf pine basal area) were used in a comparison of subplots with and without adult longleaf pine in ridge plots. In these tests, the only significant variable differing between the two sets of plots was non-longleaf pine total basal area, which doubled in subplots without longleaf pine (Table 4). In the foothill plots, the only two significantly different variables between subplots with and without longleaf pine were the species richness and non-longleaf tree total basal area variables. In a similar comparison between subplots with and without juvenile longleaf pine, four different variables (canopy openness, slope, soil depth, and tree species richness)
  • 9. 9 were significantly different between subplots. In the foothills, slope was much shallower and soil depth was much deeper than in the ridge, as seen in Table 3. The ridge subplot comparison found that canopy openness and soil depth were the two variables significantly different in subplots with and without the juveniles (Table 4). Multiple Regression Models: In the first regression model, juvenile longleaf pine frequency was the dependent variable and canopy openness, soil depth, tree species richness, slope, and non-longleaf pine basal area. The data was separated by foothills and ridge. Running a multiple linear regression test for foothill plots found that the three variables tested (slope, soil depth, and species richness, excluding non-longleaf pine basal area and canopy openness due to lack of statistical strength) were all significant in predicting juvenile frequency within the five subplots the juveniles were found in (Table 2). Analysis of the ridge plots found that soil depth, non-longleaf pine basal area, and slope did not play a significant role in predicting juvenile frequency, but canopy openness and species richness did positively correlate to the is variable significantly. The next question looked for a connection between the dependent variables of biodiversity and longleaf pine basal area and the independent variables of soil depth, slope, tree species richness, and non-longleaf pine basal area. Subplots that did not contain longleaf pine were removed and the regression was run again with the same dependent and independent variables. In the ridge plots, the significant variables in a multiple linear regression included soil depth, slope, and “other” tree total basal area. In a multiple linear regression only including these three variables, the test found a strong correlation to predicting longleaf pine basal area (R2 = 0.363, p < 0.001). Individually, slope was negatively predicted basal area, the soil depth positively predicted basal area, and the non-longleaf pine total basal area negatively predicted longleaf pine basal area (Table 2). Discussion Overall, we found that the biggest factors we tested affecting longleaf pine growth in Oak Mountain State Park were stress and competition. Stress was best measured through
  • 10. 10 two environmental stressors, slope and soil depth, competition was measured with a comparison of total basal areas of trees, and stress and competition were measured together by tree species richness. Our findings suggest that the longleaf pine have had to deal with significantly different environmental conditions within the two subsets of a montane ecosystem. The ridge is a more difficult and stressful environment for plants to grow as compared to the foothills. Also, succession due to fire suppression is more advanced in the foothills than in the ridge because invading tree species have an easier time getting a foothold there due to less stress. The ridge and foothill regions of Oak Mountain State Park appear to be very different environmentally, with significantly different slopes, soil depths, tree species richness and non-longleaf pine basal area. All of these factors seemed to have played a role in affecting the growth of juvenile and adult longleaf pine. Juvenile longleaf pine, if present, would indicate that the longleaf pines in the forest are mature enough to reproduce and the there are enough beneficial factors to allow the seeds to germinate into the juvenile stage. The differences in the variables measured must have affected regeneration of longleaf pine, seeing as regeneration nearly halted in the foothills, yet recruitment of seedlings continued on the ridge. Due to the lack of juvenile longleaf pine in the foothill subplots, the results given have to be taken as provincial findings due to small sample sizes, though this lack of juveniles in the foothills also provides evidence that juvenile longleaf pine are more likely to survive in more stressful environments due to lack of competition for space and resources. This disparity exists despite the fact that adult longleaf basal area was higher in the foothills. Adult longleaf pine basal area was used as an indirect measure of total LLP space and indicates the total space and resources taken up by this one species in a subplot. There was significantly more longleaf pine basal area in the foothill plots, meaning there was more successful growth of longleaf pine overall in the foothill plots than in the ridge plots. Environmental variables seem to directly impact growth of the longleaf pine community both on individual and collective levels. Slope was steeper in the ridge than in the foothills. Differences in slope are due to the geology of the mountain and foothills, where ridge plots are above a thick layer of sandstone, which is not as easily eroded as
  • 11. 11 the shale in the rest of the park. Sandstone prevents groundwater penetration for long term storage, hurting trees during drier periods, while steeper slopes quicken water runoff and groundwater flow. Juvenile longleaf pine were not significantly affected by the indirect effects of slope in the ridge, nor were there any differences in ridge subplots with and without juvenile longleaf pine. In the foothills, slope was negatively related to juvenile longleaf pine frequency, with significantly less slope when juvenile longleaf was present in the foothills. For adults, steeper slopes on the ridge had a negative relationship for longleaf pine basal area, though no pattern was found in the foothill plots between the two variables. Subplots with adult longleaf pine and subplots without longleaf pine did not differ for slope in either the foothills or on the ridge. Combined, these two results indicate shallower slopes generally increased longleaf pine growth, but was not the variable that allowed or prevented the establishment of the adult tree. Overall, when slope causes enough stress, it affects the tree to continue to grow, with tree growth improving on shallower slope. The data show that steeper slopes may hinder all species of tree growth, but could benefit juvenile longleaf pine due to less competition. The data also indicates that if slope was steep enough, then it could indirectly affect tree growth and therefore may affect the longleaf pine community. Soil depth was deeper in the foothills than in the ridge and may be a pivotal variable in affecting longleaf pine recruitment and growth. In foothills, juvenile longleaf pine numbers had a significant negative relationship to soil depth, but no such relationship existed in the ridge. In the foothills, soil depth was almost a perfect predictor of juvenile longleaf pine frequency and count, showing strong trends that stress in the habitat may be beneficial for longleaf pine to become established. Adult longleaf pine basal area in the ridge was marginally non-significant, but positively related to soil depth, with deeper soils indicating greater basal area, while there was no significant trend found in the foothills. There was also no significant difference in subplots where adult longleaf pine was present or absent in either the foothills or the ridge. The lack of change between subplots with and without adult longleaf pine indicates that soil depth was not the variable that allowed or prevented the establishment of the adult tree. When compiling all of this data, soil depth provides evidence that it causes stress that affects longleaf pines’ continued growth when shallow enough and that the deeper the soil, the easier for the tree to grow. Like with slope, if soil depth was not
  • 12. 12 extreme enough, then it would not play a direct factor in the longleaf pine community. Finally, longleaf pine generally grows better in deeper soils, but the juvenile longleaf pine have more success in shallower soils due to less competition. The shallower soil may be a limiting factor in survival for other plants, but not for juvenile longleaf pine; this opens up space and resources for more juvenile longleaf pine to germinate and grow. Non-longleaf pine basal area, being greater in the foothills than the in the ridge, seemed to have a negative impact on the ability for adult longleaf pine to grow. Non-longleaf pine basal area was used as a way to measure the amount of space non-longleaf pine trees took in the two regions of the park, giving an idea of the total area of trees are found in each subplot. There was significantly more tree area in the foothills than as compared to the ridge, supporting that the notion that higher amounts of stress lead to lower amounts of total tree growth. Overall, the only significant regression result for both juvenile and adult growth was for adult longleaf pine basal area in the ridge, which has a negative relationship. There was also a significant drop in non-longleaf pine basal area when longleaf pine were in the area. This supports the idea that other trees generally compete against and are bad for the LLP community. With more stress, there will be both lower non-longleaf pine basal area and higher longleaf pine take up space per subplot. Combined with tree species richness results, the data may provide evidence that that non-longleaf pine trees and the longleaf pine community have an antagonistic relationship. Tree species richness was also greater in the foothills than in the ridge. Tree species richness was used as an indicator of the state of the ecosystem, with increased richness meaning greater competition for longleaf pine trees. This variable also pointed to possible stress variations between ridge and foothills. In the ridge, tree species richness was negatively related to longleaf pine basal area, meaning increased diversity related to the declining health of longleaf pine community, while decrease in diversity would be beneficial to longleaf pine trees. Tree species richness was greater in foothill subplots without adult longleaf pine, supporting the idea that as species richness increases, the more competition and the worse it is for the longleaf pine community. The ridge plots were not significantly affected by species richness, which due to the lesser amounts competition from increased stress. For juvenile longleaf pine, tree species richness was positively related in the ridge, but negatively related in the foothills. The mixed results support the idea that lower levels of
  • 13. 13 stress in a high stress environment create higher chances of germination and growth into the juvenile stages. In the foothills, more stress is needed to weed out competition for juvenile longleaf pine to grow, so more stress means less competition. The findings suggest that longleaf pine compete with other tree species for resources: the more resources the longleaf pine obtain, the more area they are able to take. Juvenile longleaf also have to compete with other species when there is little stress, but when stress is elevated to affect the amount of species, juvenile longleaf pine only need to worry about stressors like soil depth and slope. We were very surprised to find that canopy openness lacked any significant trends with any of the tests performed. Young longleaf trees need many hours of direct sunlight to survive and grow each day. Canopy openness is directly related to light availability at the forest floor. When more light allowed to reach the forest floor, undergrowth receive more energy to photosynthesize and produce sugars to store, making it an important factor in small plant growth. There was one marginally non-significant result when comparing canopy openness with juvenile longleaf pine in the ridge, with a positive relationship between the two. The number of insignificant results provide evidence that canopy openness did not play a major role in longleaf pine recruitment or growth. This lack of difference may exist because both areas may be at or close to full closure, which is especially true in the foothills with an increased invasion of broadleaf trees. Conclusion: Stress is helpful for LLP as steeper slopes and shallower soils introduce stress that has been waning with fire suppression. The lack of stress has allowed increased competition against the longleaf pine, with elevated tree species richness and growth of non-longleaf pine negatively and unevenly impacting this community the foothills region of the park. These have seemed to increased stress in the longleaf pine community to have a greater negative impact than slope and soil depth do. Both of these could be key factors in successful germination, with soil depth being significantly lower and slope being significantly greater when comparing plots with and without juvenile LLP. Using these initial findings, we can start looking into how these variables predict and/or affect the ability for this tree to establish itself from the juvenile stage into an adult and how it continues to grow throughout its lifetime.
  • 14. 14 In future research, other factors that should be looked into include temperature, slope aspect, and the average distance away from the parent tree. All of these could also be factors that cause changes in the abundance of juvenile longleaf pine and size of adult LLP in each of the subplots. For example, ridge slopes face southeast, meaning the summer sun is more intense on the slope than others facing different directions, drying the soils more quickly than on the north facing slopes. How does this change with different slope aspects? Chance also may play a large factor of deciding whether a juvenile or adult longleaf pine will grow from a seed into a juveniles or juveniles into an adult, respectively. Using these initial findings, we can start looking into how these variables predict and/or affect the ability for this tree to germinate and grow from seeds into the juvenile stage and into an adult. There is generally poor recruitment of longleaf pine in the park overall, so how can that be improved upon by the trees? With this poor recruitment means a potential drop in adult longleaf pine in the future. With this initial research, we will need to look for more a detailed understanding of what other environmental factors play a role in the growth of longleaf pine to help them maintain and re-extend their community in the future.
  • 15. 15 Tables and Graphs Table 1. Resultsof a Mann-WhitneyUtestseeingif there were differencesbetweenthe ridge and foothill regionsin OakMountainState Parkin Pelham, AL. A p value of lessthan.05 indicatesa significantdifference. Foothill Ridge Variable Mean (SD) Range Mean (SD) Range P value Canopy Openness (%) 12.7 (1.7) 7.9 – 17.5 14.5 (5.5) 9.5 – 36.7 0.176 Slope (⁰) 12.0 (5.8) 1.0 – 28.0 21.0 (6.0) 5.0 – 41.0 <0.001 Avg Soil Depth (cm) 34.2 (13.9) 4.9– 65.2 18.3 (13.9) 0.0 – 53.0 <0.001 Juvenile LLP Count 2.8 (1.8) 1.0 – 5.0 12.2 (16.3) 1.0 – 53.0 <0.001 LLP BasalArea (m2 ) 0.18 (.15) 0.0024 - 0.84 0.11 (0.09) 0.0005 -0.47 0.004 Tree Species Richness 5.6 (2.0) 2.0 – 11.0 4.1 (1.5) 1.0 - 9.0 <0.001 Non-LLP BasalArea (m2 ) 0.13 (.09) 0.016 - .51 0.065 (.06) 0.0009 - 0.33 <0.001 Table 2. Resultsof multiple linearregressionsof the numberof juvenile longleaf pine oradultlongleaf pine basal area(dependentvariables) againstCanopyOpenness(and/orSpeciesrichness),Slope,basal area of Non-longleaf trees,andSoil Depth(independentvariables) intwodifferent topographicregions of OakMountainState Parkin Pelham,AL.Pvalueslessthan.05 representsignificantR2 values. Region Model’sDependent variable R2 P Independent variables Beta P Foothills JuvenileFrequency 1.000 0.011 Slope -0.442 0.020 Soil Depth -0.953 0.006 SpeciesRichness -0.232 0.038 AdultBasal Area 0.204 0.030 CanopyOpenness 0.224 0.124 Slope 0.144 0.281 Soil Depth -0.105 0.438 SpeciesRichness -0.214 0.139 Non-LLPBasal Area -0.132 0.393 Ridge JuvenileFrequency 0.355 0.001 CanopyOpenness 0.313 0.067 Slope -0.274 0.145 Soil Depth 0.065 0.845 SpeciesRichness 0.537 <0.001 Non-LLPBasal Area -0.105 0.431 AdultBasal Area 0.387 <0.001 CanopyOpenness -0.176 0.115 Slope -0.386 0.001 Soil Depth 0.208 0.068 SpeciesRichness -0.024 0.824 Non-LLPBasal Area -0.280 0.014 Table 3. Comparisonof subplotswithandwithoutjuvenile longleaf pineinOakMountainState Parkin Pelham,AL.Pvalueslessthan.05 representasignificant difference betweenthe subplotsusingaMann- WhitneyU test. With Juvenile LLP Without Juvenile LLP Region Variables Mean (SD) Range Mean (SD) Range P Value Foothills CanopyOpenness(%) 12.5 (1.7) 10.2 – 14.0 13.9 (5.0) 9.6 – 32.8 0.447
  • 16. 16 Slope (⁰) 12.6 (5.8) 7.0 – 28.0 19.9 (5.7) 7.0 – 32.0 0.002 AvgSoil Depth(cm) 32.52 (14.0) 15.7 – 54.2 23.9 (13.3) 0.0 – 48.1 0.002 SpeciesRichness 4.8 (2.0) 2.0 – 7.0 5.8 (1.9) 3.0 – 11.0 0.437 Ridge CanopyOpenness(%) 14.4 (5.5) 9.8 – 36.7 13.9 (5.0) 9.6 – 32.8 0.466 Slope (⁰) 21.5 (6.0) 9.0 - 38.0 19.9 (5.7) 7.0 – 31.5 0.357 AvgSoil Depth(cm) 16.8 (13.7) 0.0 – 48.4 23.9 (13.3) 0.0 – 48.1 0.103 SpeciesRichness 3.7 (1.3) 2.0 – 6.0 4.3 (1.4) 1.0 – 9.0 0.155 Table 4. Comparisonof subplotswith andwithoutadultlongleaf pineinOakMountainState Parkin Pelham,AL.Pvalueslessthan.05 representasignificantdifference betweenthe subplotsusingaMann- WhitneyU test. With adult LLP Without adult LLP Region Variables Mean (SD) Range Mean (SD) Range P Value Foothills Slope (⁰) 12.5 (5.7) 2.0 – 28.0 11.4 (5.8) 1.0 – 26.0 0.423 AvgSoil Depth(cm) 34.1 (13.8) 4.9 – 64.4 34.2 (14.4) 12.3 – 65.2 0.908 SpeciesRichness 5.6 (1.9) 2.0 – 11.0 5.73 (1.9) 3.0 – 11.0 <0.001 Non-LLPBasal Area(m2 ) 0.13 (0.09) 0.016 – 0.51 .20 (0.09) 0.070 - 0.44 <0.001 Ridge Slope (⁰) 21.0 (6.0) 5.0 – 41.0 21.1 (5.8) 8.0 – 32.0 0.886 AvgSoil Depth(cm) 17.8 (13.8) 0.0 – 48.4 19.5 (13.3) 0.0 – 48.1 0.564 SpeciesRichness 4.14 (1.4) 1.0 – 9.0 3.1 (1.26) 2.0 – 5.0 0.966 Non-LLPBasal Area(m2 ) 0.065 (0.063) 0.0 – 0.22 0.11 (0.064) 0.017 - 0.33 <0.001
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