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Observations on a Rare Old-Growth Montane Longleaf Pine Forest in Central
North Carolina, USA
Author(s): Thomas W. Patterson and Paul A. Knapp
Source: Natural Areas Journal, 36(2):153-161.
Published By: Natural Areas Association
DOI: http://dx.doi.org/10.3375/043.036.0206
URL: http://www.bioone.org/doi/full/10.3375/043.036.0206
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Volume 36 (2), 2016 Natural Areas Journal 153
ABSTRACT: Montane longleaf pine (Pinus palustris) forests are rare and no detailed inventory exists
documenting stands in North Carolina. We inventoried all longleaf pine trees (n = 403) growing in
a 24-ha remnant montane longleaf pine forest in the Uwharrie Mountains of central North Carolina,
USA, in autumn 2014 to (1) map their location, (2) document age/height/diameter characteristics, and
(3) determine special ecological features of this rare montane population. All longleaf pine were geo-
graphically referenced via GPS, measured for height and diameter, and a subsample of trees was cored
to determine age. All longleaf pine were mapped based on growth-stage categories—grass, juvenile,
young adult, and mature—to determine spatial patterning of stand-age characteristics. The longleaf
pine stand contains a variety of growth-stage categories, but is dominated (63%) by mature-stage trees
growing on south- and southwestern-facing slopes, while nearly all regeneration-stage trees (i.e., grass
and juvenile) are growing on northwest-facing slopes, suggesting environmental conditions conducive
to establishment have changed. Median (maximum) tree height and trunk diameter for young adult and
mature were 17 (25) m and 38 (72) cm, respectively. Median (maximum) tree age at 0.3 m height was
116 years (272 years), and at least seven trees were greater than 150 years old, with four trees establishing
in the 18th century. We conclude that the stand’s characteristics—400+ trees of various ages including
old-growth, occurring principally on steep, southerly slopes with a total relief of 85 m, and extending
over 24 ha—warrant “montane” longleaf pine forest status in North Carolina.
Index terms: forest inventory, longleaf pine, montane, North Carolina, old growth
INTRODUCTION
Longleaf pine (Pinus palustris Mill.) is
a long-lived (>400 years; Earle 2014)
species native to the southeastern United
States Atlantic and Gulf Coastal Plains
and Piedmont physiographic regions,
but also occurs at some locations in the
Ridge and Valley (i.e., montane) regions
of Alabama and Northern Georgia (Boyer
1990). Since the late 17th century, the
species’ extent has diminished from 37
million hectares (Frost 1993) to 1.74 mil-
lion hectares (Oswalt et al. 2012) due to
various anthropogenic processes including
fire suppression, logging, agriculture, and
naval stores activities (Frost 2006). Within
the Piedmont physiographic region ranging
from Alabama to North Carolina, some
longleaf pine communities exist on steep,
mountainous terrain at elevations typically
<610 m above sea level and represent the
most atypical longleaf pine forest com-
munity type classified (Peet 2006). These
stands, often called “montane,” are most
common on southerly and southwesterly
slopes, with trees typically growing on
well-drained, rocky soils, and occur in
mixed pine (e.g., P. echinata Mill., P. taeda
L., and P. virginiana Mill.) and hardwood
(e.g., Quercus marilandica Münchh, Q.
stellata Wangenh) mesophytic forests
(Harper 1905; Maceina et al. 2000; Peet
2006). Here, we discuss an ecologically
rare and newly managed montane longleaf
pine forest in the Uwharrie Mountains of
central North Carolina.
Documentation regarding montane longleaf
pine communities is sparse, in part due to
the paucity of extant stands and, thus, our
understanding about these rare forests is
limited. Vegetation surveys (Maceina et al.
2000) and stand dynamics studies (Varner
et al. 2003; Stokes et al. 2010) have been
undertaken at montane longleaf pine stands
located within the Mountain Longleaf
National Wildlife Refuge near Anniston,
Alabama. Similarly, response to manage-
ment changes for both bird communities
(Kronenberger et al. 2014) and herbaceous
plants and grasses (Cipollini et al. 2012)
was studied at a remnant montane longleaf
pine stand owned by Berry College in Floyd
County, Georgia. A number of conference
proceedings, reports, and graduate theses
also exist that have documented various
aspects of these montane longleaf pine
forests, but again, with a focus on stands
in Alabama and Georgia.
To our knowledge, no formal inventory
regarding North Carolina’s mountainous
longleaf pine communities has occurred.
Wells (1974) mentions a stand of “about
20 canopy-height” longleaf pine with the
largest diameter at breast height (DBH)
values of 45 cm at an unspecified location in
the Uwharrie Mountains. Similarly, Bates
(2001) identifies a 17-ha “remnant” stand
of longleaf pine on “an unnamed mountain
north of Daniel Mountain” in the Uwhar-
Natural Areas Journal 36:153–161
R E S E A R C H A R T I C L E
2 Corresponding author: twpatter@uncg.
edu; (336) 334-5388
•
Observations on a
Rare Old-Growth
Montane Longleaf
Pine Forest in
Central North
Carolina, USA
Thomas W. Patterson1,2
1Carolina Tree-Ring Science Laboratory
Department of Geography
University of North Carolina Greensboro
Greensboro, NC 27402-6170
Paul A. Knapp1
•
154 Natural Areas Journal Volume 36 (2), 2016
rie National Forest (“Gold Mine Branch
Longleaf Pine Forest”), but did not provide
age/growth specifics. This latter longleaf
pine forest (hereafter known as Gold Mine
Branch) is well preserved and contains
seedling through old-growth individuals
approaching 300 years of age. Here we
discuss our observations following an in-
ventory of all longleaf pine trees within the
Gold Mine Branch forest. Specifically, we
report on the longleaf pine trees in terms of
(1) the geographic patterns of occurrence,
(2) age/height/diameter characteristics, and
(3) special features of this rare mountain-
ous population.
METHODS
Study Site
Gold Mine Branch Longleaf Pine Forest is
located in the 205 km2 Uwharrie National
Forest at 35.416◦ N, 80.036◦ W (Figure 1),
1 km west of the Uwharrie River and 14 km
northwest of the town of Troy, North Caro-
lina. Uwharrie Mountain geology consists
of weathered, Cambrian era interbedded
felsic and mafic metavolcanics extending
46 km between Randolph, Montgomery,
and Stanley counties (Rogers 1982; Daniel
and Butler 1996). These mountains range
from 150 to 335 m elevation, have up to
150 m of topographic prominence, and
support Piedmont and disjunct popula-
tions of mountain and coastal plain species
(Wells 1974; Bates 2001). The site was
identified in Bates’ (2001) Montgomery
County Natural Heritage Inventory as a
17-ha remnant Piedmont-longleaf forest.
Bates (2001) noted the presence of longleaf
pine on the steep south- to southwest-facing
slopes of an unnamed mountain north of
Daniel Mountain that is separated by the
perennial stream, Gold-Mine Branch. A
rocky substrate of Georgeville silt loam
dominates the site (Soil Survey Staff 2015),
Figure 1. Location of all live longleaf pine trees by size class at Gold Mine Branch in the Uwharrie National Forest. Size classes are as follows: (1) grass
stage, (2) juvenile/bottlebrush (i.e., individuals without lateral branch growth), (3) young adult (lateral branches present and DBH ≥10 cm but ≤20 cm, and
(4) mature (>20 cm DBH). Contour interval is 3 m.
Volume 36 (2), 2016 Natural Areas Journal 155
and slopes approach 25○ (Rogers 1982).
Elevations at the top (266 m) and bottom
(134 m) of the Gold Mine Branch site rep-
resent 132 m of total vertical relief. Mean
annual precipitation for the region is 120
cm and is distributed evenly throughout
the year. Mean July (January) highs aver-
age 31 oC (10 oC), with maximum and
minimum temperatures of 39 oC and -9
oC (US Climate Data 2015 ).
Gold Mine Branch was purchased in 1944
from the Russell Family, who owned land in
the region for timber and mineral extraction
(D. Walker, US Forest Service, pers. comm.
2015). From 1944 to 1961, the land was
managed by the North Carolina Wildlife
Resource Commission, primarily for hunt-
ing and recreation, and thereafter by the
US Forest Service for similar purposes. The
site was listed as a Special Interest Man-
agement Area in 2012 for its botanic and
scenic characteristics (US Forest Service
2012). Prescribed fires were set in 2010
and 2013, and two wildfires are thought to
have occurred at the site in the late 1970s
and 1920s (D. Walker, pers. comm.). Nu-
merous (estimated 150–200) pine stumps
exist throughout the stand that were likely
cut during the early 20th century and
represent a significant disturbance event,
but we saw no evidence of previous naval
stores activity (i.e., box cuts or chevrons)
on either the stumps or live trees. Fire
scars occurred on approximately 7% of
the mature trees and almost exclusively
on the upslope side of the boles.
The canopy at Gold Mine Branch consists
of longleaf and shortleaf pine, along with
chestnut (Quercus montana Willd.), post,
and blackjack oaks. Other canopy species
include red maple (Acer rubrum L.) along
with mockernut (Carya tomentosa Lam.)
and pignut (C. glabra Mill.) hickories.
Understory vegetation includes muscadine
grape (Vitis rotundifolia Michx.), asters
(Pitysopsis graminifolia (Michx.) Nutt.,
Coreopsis verticillata L., Cunila origanoi-
des L.), bracken fern (Pteridium aquilinum
(L.) Kuhn), Goat’s rue (Tephrosia virgin-
iana (L.) Pers.), shrub species including
St. Andrew’s cross (Hypericum hypericoi-
des (L.) Crantz subsp. hypericoides), St.
John’s-wort (H. stragalum L.), Deerberry
(Vaccinium stamineum L.), Sparkleberry
(V. arboretum Marshall), and Horse sugar
(Symplocos tinctoria (L.) L’Hér.), and the
significantly rare Piedmont indigo bush
(Amorpha schwerinii C.K. Schneid.)
(Bates 2001).
Sampling Procedures
Field data were collected during Novem-
ber and December, 2014. Following an
initial site survey, we determined that the
Gold Mine Branch longleaf pine commu-
nity was characterized by higher-density
“core” and lower-density (often a single
tree) “periphery” populations and required
different sampling methods to document.
Within the core population, where multiple
trees were within 15 m of each other, we
marked 50-m-wide transects with flagging
and documented all live trees along a
vertical gradient. For live trees located in
the periphery (Figure 1), we walked along
east-west-running constant-elevation gradi-
ents from the westernmost to easternmost
sections of Gold Mine Branch at 50-m
elevation gradients, again marking intervals
with flagging. For each tree, we recorded
location using a Garmin 60CSx GPS unit
and classified individuals as follows: (1)
grass stage, (2) juvenile/bottlebrush (i.e.,
individuals without lateral branch growth),
(3) young adult (lateral branches present
and DBH ≥10 cm but ≤20 cm), and (4)
mature (>20 cm DBH).
All young adult and mature trees were
measured for DBH using diameter tape,
and canopy height was measured using a
Nikon Forestry 550 laser range finder. We
removed one increment core sample each
from 50 young adult and mature trees of
various diameters at approximately 0.3-m
height using a 5.15-mm Haglof increment
borer to determine tree age. Trees were
selected to represent a range of DBH
values and included all trees with visual
old-growth characteristics (smooth bark,
twisted boles, sparse and flat-topped crown
(Pederson 2010)) without external evidence
of heart rot. These tree-ring data were then
used to develop a predictive DBH/age
model for the remaining trees sampled.
Laboratory Procedures
We entered all longleaf pine locations in
ESRI ArcGIS 10.2 (ESRI 2013) for inven-
tory, mapping, and geospatial analysis.
Tree cores were air-dried and mounted
to wooden strips with wood glue, then
sanded with progressively finer sandpaper
(120, 220, 400 grit) until all annual growth
rings were visible. Age determination was
made by crossdating using the list method
(Yamaguchi 1991) for all cores dating to the
1850s. Tree-age at 0.3 m for samples dat-
ing pre-1850 was estimated by ring count
as the lack of cores prevented crossdating.
Thus, dating through 1850 is marked by
annual precision, whereas interior dates of
older trees may be imprecise because of
possible missing or false rings.
Upon core processing we determined that
approximately 14% of the sampled trees
contained heart rot, therefore a true age at
0.3 m could not be achieved as the interior
of the tree, including the pith, was absent.
These cores were removed from further
age/DBH analysis. Additionally, longleaf
pine can remain in the grass stage for up
to 20 years, all the while not producing
annual growth-rings (Pessin 1934). Thus,
our predictive DBH/age model produced
relative, as opposed to absolute, tree ages.
We used tree age at 0.3-m height from the
43 remaining cores to build the following
predictive model from trunk diameter:
Predicted tree age at 0.3 meters = β0 +
β1 DBH
Where β0 is the intercept of the regression
line and β1 dictates the slope. We could then
assign a predicted tree age at 0.3 m for the
remaining 245 young adult and mature
longleaf pine that were not cored.
RESULTS AND DISCUSSION
Sample Size and Site Expansion
Our study identified 403 longleaf pines
growing at Gold Mine Branch, including
41 grass stage, 67 juvenile stage, 39 young
adult, and 256 mature trees. During our
data collection, we identified an additional
156 Natural Areas Journal Volume 36 (2), 2016
south-to-east-facing slope undocumented
in Bates’ survey (2001) that expanded the
site from 17 to 24 ha. The lowest elevation
tree was at 170 m and the highest was at
255 m, giving a total relief of the longleaf
stand of 85 m.
Height, Diameter, and Location
Median (maximum) tree height and trunk
diameter for all mature trees were 17 (25) m
and 38 (72) cm, respectively (Figure 2a and
b). Tree density ranged from 20 to 60/ha,
with a basal area of 2.2–4.5 m2/ha from
periphery to core areas. Approximately
one third of all longleaf pine at Gold Mine
Branch were growing on southwesterly
slopes, followed by 22% on northwesterly
slopes. No trees were located on northeast-
facing slopes. Only five trees were on
north slopes, and three trees were on east
slopes, cumulatively accounting for 2%
of all longleaf pine. We assessed whether
longleaf pine in the four stages of growth
were occurring throughout the site or were
limited to specific aspects. The majority
of longleaf pines in the grass and juvenile
stages were on northwest-facing slopes.
More young adult trees were on northwest
and southwest slopes, and mature trees
were predominantly found on southwest
and southeast slopes (Figure 3).
Relative Age
The predictive DBH/age model accounted
for approximately 27% (r2 = 0.265, P <
0.001) of age variance (Figure 4). The
model overestimated the youngest trees
by up to 40 years and underestimated the
oldest trees by over 100 years, suggesting
that DBH is a weak predictor of longleaf
pine age at Gold Mine Branch. Thus, our
ability to accurately assess relative age
based on the predictive age model is limited
and we refrain from further assessment
based on predicted age. Of the 50 samples
collected, only 43 trees were datable to
pith. Mean (median) relative age at 0.3
m was 116 (102), with seven (five) trees
exceeding 150 (200) years old (Figure 2c).
The age gap between the five oldest trees
dated >190 years and the next-oldest tree
dated at 159 years suggests nearly all trees
Figure 2. Histograms of frequency for (a) height and (b) trunk diameter for all young adult and ma-
ture trees (n = 295). Minimum age (c) for all young adult and mature trees based on a subsample of
43 trees.
Volume 36 (2), 2016 Natural Areas Journal 157
greater than approximately 50–60 years
old in the early 1900s were logged. Of the
documented old trees remaining, all had
marked bole deformations (e.g., twisted,
snapped top, leaning), suggesting that they
may have been of lesser harvestable value.
Additionally, at least two other longleaf
pine at the site have similar hallmarks
of multicentury age (>200 years, Figure
5a), but we were unable to obtain tree age
because of heart rot.
Special Features of Gold Mine Branch
(1) The oldest tree dated to pith at AD
1743 at 0.3 m height, and we estimated
its age to be over 270 years old. Similarly,
the next oldest tree had an innermost ring
that dated to 1761 at 0.3 m height before
heart rot. Accounting for heart rot and a
Figure 3. Compass plots of longleaf pine in the four growth stages. Note difference in scales for the number of trees on the vertical axis.
grass stage without ring development, we
estimate both trees could approach or ex-
ceed 300 years old. Based on the number
of longleaf pines >150 years old (Figure
2c), and the abundance of visually gnarled
hardwoods that co-occur throughout the
site, Gold Mine Branch is a rare, old-growth
montane longleaf forest and unique in areal
extent for North Carolina. Additionally,
throughout the site we identified remnant
longleaf pine stumps that were likely felled
before a time (pre-1950) of mechanized
saws during the Russell Family ownership.
These meter-high stumps are rich in resin
and susceptible to fire if not preserved. The
abundance of longleaf pine in the 80–120
age range, coupled with a presence of old
stumps, suggests an early 20th century
felling operation likely influenced by the
construction of railways in Montgomery
County in the 1890s for timber extraction
(Carriker 1992).
(2) The majority of grass-stage and juvenile
trees at Gold Mine Branch are growing on
northwest-facing slopes, while the majority
of young adult and mature trees are grow-
ing on southwest-facing slopes (Figures 3
and 5b). Why the distinction exists is un-
known, but suggests competitive exclusion,
changing environmental conditions, or a
combination of these has occurred since
the area was logged. The 2010 and 2013
fires may have contributed to the observed
spatial pattern as wildfires tend to burn hot-
ter on south-facing slopes (Heyerdahl et al.
2001), and juvenile longleaf fire mortality
(up to 80% ) is greater in mature stands due
to litter accumulation, resulting in hotter
burning fires (Grace and Platt 1995).
158 Natural Areas Journal Volume 36 (2), 2016
(3) Gold Mine Branch topography—locat-
ed along a 1.5-km-length east-west oriented
ridgeline with southerly aspect slopes—is
unmatched in areal extent in the Uwharrie
Mountains and provides extensive steep
terrain suitable for longleaf pine establish-
Figure 4. Scatterplots of the regression line of the predicted DBH/age model (top) and height and trunk diameter for all young adult and mature trees (bot-
tom).
ment. We posit that the extensiveness of the
montane longleaf pine forest at this location
is a result of the rare environmental condi-
tions in the Uwharrie Mountains that confer
a competitive advantage to this species in
an environment otherwise dominated by
other pine and hardwood species. Further,
the steep, boulder-strewn terrain may have
hindered the logging of all the trees, leav-
ing behind a population base sufficient for
species propagation.
Volume 36 (2), 2016 Natural Areas Journal 159
(4) A 2-ha stand of scattered montane
longleaf pine is located approximately 8 km
south of Gold Mine Branch (D. Walker, US
Forest Service, pers. comm. 2015). While
this remnant forest near Dennis Mountain
matches the description by Wells (1974)
of nearly two dozen mature longleaf pine
trees growing on south- and southwest-
Figure 5. (a) One of the authors posing with a “bonsai topped” old-growth longleaf pine, and (b) two juvenile and one young adult longleaf pine growing on
the northeast slope. All cored bonsai trees had interior dates in the 1700s.
facing slopes between 198 and 229 m,
we confirmed (Julie Moore, US Fish and
Wildlife Service, pers. comm. 2015) that
Wells (1974) was, in fact, writing about
Gold Mine Branch. No other stands of
montane longleaf pine forest exist in the
Uwharrie Mountains that match the spatial
and topographical extent of Gold Mine
Branch.
CONCLUSIONS
We suggest that Gold Mine Branch should
be classified as a montane longleaf pine
forest. Bates (2001) identifies the stand
as a “Piedmont Longleaf Pine Forest”
—nomenclature consistent with the North
Carolina Natural Heritage Program (Scha-
160 Natural Areas Journal Volume 36 (2), 2016
fale 2012) that includes inland longleaf
pine forests of the undulating topography
of central North Carolina (Moore and
Fogo, a paper presented in the 3rd Mon-
tane Longleaf Pine Conference, Auburn,
Alabama, 2008). That said, montane is
geographically descriptive and several
significant environmental differences war-
ranting distinction exist between Gold
Mine Branch and the surrounding Pied-
mont longleaf pine forests. The Uwharrie
Mountains are classified as a physiographic
subprovince of the Piedmont region marked
by “sharp relief” (Hack 1982). The steep
and principally south-facing terrain of Gold
Mine Branch likely produces a different
temperature regime than flatter landscapes,
with significant differences in maximum
daily temperatures based on aspect and
daily minimum temperatures related to
slope position (Bolstad et al. 1998). Rates
of evapotranspiration are higher on south-
facing slopes than flatter terrain (Holland
and Steyn 1975; Countryman 1978), and
soil-moisture regimes are impacted (e.g.,
Countryman 1978). Fire regimes in moun-
tainous areas are different as fires travel
faster upslope (Butler et al. 2007) and may
be hotter on south- versus north-facing
slopes (Rothermel 1972; Heyerdahl et al.
2001). Finally, trapped forest duff and litter
on the upslope side of trees allows fires to
burn hotter than elsewhere (Arno and Sneck
1977) and is consistent with the location
of fire scars found almost exclusively on
the upslope side of mature longleaf pines
at Gold Mine Branch.
Our inventory recorded most and, perhaps,
all the live longleaf pines at Gold Mine
Branch, excluding seedlings, and this
documentation provides benchmark data
for the Uwharrie National Forest Service
and future ecological studies at the site. The
2012 Uwharrie National Forest Manage-
ment Plan places emphasis on reforesting
longleaf pine throughout the forest and has
placed Gold Mine Branch under Special
Interest Management, where limitations
to thinning and burning are prescribed to
enhance the site’s properties (US Forest
Service 2012). It is our hope that these
findings may help in better understand-
ing the effects of ongoing management
practices at Gold Mine Branch.
ACKNOWLEDGMENTS
We are grateful for the assistance of many
individuals who offered insights, field
assistance, and technical support. Specifi-
cally, we thank Nell Allen, Moni Bates,
Boon Chesson, Laura Fogo, Doug Gold-
man, Anna Levi, Julie Moore, Selima Sul-
tana, Deborah Walker, and Keith Watkins.
Additionally, we thank two anonymous
reviewers for their help in improving this
manuscript. This research was supported in
part by the National Science Foundation,
Grant No. DGE0947982.
Thomas Patterson is a PhD candidate in
the Carolina Tree-Ring Science Labora-
tory, Department of Geography at the Uni-
versity of North Carolina at Greensboro.
His research interests have focused on
dendroecological applications in longleaf
pine forests.
Paul Knapp is a professor and director of
the Carolina Tree-Ring Science Laboratory
in the Department of Geography at the Uni-
versity of North Carolina at Greensboro.
His interests have principally focused on
the paleoecological dynamics of western
North American forests.
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