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2006 SOUTHEASTERN NATURALIST 5(4):637–648
Summer Meleagris gallopavo silvestris Use of a Landscape
Dominated by Agriculture and Pinus spp. Plantations
John J. Morgan1,*, Sara H. Schweitzer2, and John P. Carroll2
Abstract - Meleagris gallopavo silvestris (Eastern Wild Turkey) habitat was altered
in the Southeast by the introduction of Pinus spp. plantations to agricultural areas
through the Conservation Reserve Program. However, the preponderance of M.
gallopavo silvestris research has focused on extensive Pinus spp. plantations that
lack the cover-type diversity that typifies the Southeast. From May–July 1998 and
1999, we monitored 36 radio-tagged M. gallopavo silvestris in Burke County, GA to
investigate habitat use in landscapes intensively managed for agriculture and silviculture.
We used compositional analysis to identify habitats selected by male and
female M. gallopavo silvestris during summers. Proportions of habitat types within
the home range were different from habitats at radio-locations of males and females.
Hardwood stands and fields were the most-selected habitat types by M. gallopavo
silvestris in the summer. However, within home ranges, males and females also
selected closed-canopy Pinus spp. habitats. Hens with broods did not preferentially
select planted Pinus spp. habitats, but their use of Pinus spp. stands was greater than
use of agricultural fields. The replacement of agricultural fields by closed-canopy
Pinus spp. plantations may have improved habitat quality for M. gallopavo silvestris
in some areas of the Southeast by diversifying the landscape. Our results suggest that
closed-canopy planted Pinus spp. cover types are not detrimental to M. gallopavo
silvestris when well distributed with fields and mature hardwood drains.
Periodically since the 1930s, the United States Department of Agriculture
(USDA) has introduced conservation programs to reduce agricultural soil
erosion. The Conservation Reserve Program (CRP), a provision within the
Food Security Act of 1985, was adopted to control soil erosion and curb
agricultural commodity surpluses. The CRP subsidized landowners over a 10-
or 15-year period in exchange for establishment of permanent vegetation on
In the Southeast, more than 800,000 ha were planted in Pinus spp. (pine),
which comprised the majority of CRP contracts (Farm Service Agency
1997). Once these Pinus spp. plantations reached canopy closure, their value
to some wildlife decreased (Allen 1993, Felix et al. 1986). Many wildlife
biologists thought that CRP practices and public funds should benefit wildlife,
not create Pinus spp. plantations (Allen et al. 1996). The 1996 Farm Bill
introduced the Environmental Benefits Index (EBI) and listed wildlife as a
co-equal to soil and water quality. Applications were ranked by EBI score,
1One Sportsman’s Lane, Frankfort, KY 40601. 2Warnell School of Forestry and
Natural Resources, University of Georgia, Athens, GA 30602. *Corresponding
author - email@example.com.
638 Southeastern Naturalist Vol. 5, No. 4
so CRP funds maximized public gains with respect to wildlife habitat, soil
erosion, and water quality. The score improved if established Pinus spp.
plantations were thinned.
Meleagris gallopavo silvestris Vieillot (Eastern Wild Turkey) use of
pine-dominated habitats across the Southeast has been well documented.
The study area of Exum et al. (1987) in Alabama was 80% Pinus spp. and
supported a M. gallopavo silvestris population that was hunted each spring.
In other studies, hens tended to prefer Pinus spp. plantations in the
spring and summer (Burk et al. 1990, Palmer et al. 1993), and males used
planted Pinus spp. in the summer (Exum et al. 1987, Wigley et al. 1985).
However, we were uncertain how M. gallopavo silvestris use habitats within
landscapes fragmented by Pinus spp. plantations and agricultural fields that,
in many cases, resulted from the CRP.
We investigated habitat use and home ranges of M. gallopavo silvestris
within a fragmented landscape dominated by agricultural areas and Pinus spp.
plantations. We categorized Pinus spp. plantations as open- (herbaceous
understories) or closed-canopy (barren understories) to determine if stand
structure influenced M. gallopavo silvestris habitat use. We investigated M.
gallopavo silvestris habitat use within our study area and within home ranges.
Our study originated at Di-Lane Plantation Wildlife Management Area
(WMA), Upper Coastal Plain physiographic region, 1.6 km south of
Waynesboro, GA. Di-Lane Plantation WMA is a 3278-ha area managed by
the Georgia Department of Natural Resources (GADNR) for wildlife since
1992. Previous owners farmed the land for row crops through 1991, and in
1988, they enrolled 286 ha in CRP by planting Pinus taeda (L.) (loblolly
pine) as the cover crop (CP3) at 1793 seedlings/ha. Management of the
WMA by GADNR focused on Colinus virginianus L. (Northern Bobwhite)
and included prescribed burning of fields and upland hardwoods (generally
late winter by aerial or hand ignition), seasonal disking, and supplemental
food plantings (annual grains, Trifolium spp. [clover], and cool-season grass
plots). Non-CRP Pinus spp. stands were row-thinned by 50% in 1996 followed
by annual disking of thinned rows to promote annual weeds and bare
ground for C. virginianus. In spring 1997, a 15-ha CRP stand was rowthinned
(33%), and a 17-ha CRP stand was strip-thinned (40%) and
row-thinned (33%) for research purposes. Open-canopy planted Pinus spp.
(young stands < 8 years old and thinned stands 12–15 years old) were
typified by understories of assorted annual weeds including agricultural
pests like Senna obtusifolia Irwin and Barneby (coffeeweed) and Sorghum
halepense Moench (Johnson grass), whereas closed-canopy Pinus spp. (8–
20 years old) were comprised of needle-laden ground cover nearly devoid of
herbaceous plants (S.H. Schweitzer, unpubl. data). Most CRP Pinus spp.
plantations were adjacent to mature hardwood stands and in some instances
bordered row crops or fields as well.
2006 J.J. Morgan, S.H. Schweitzer, and J.P. Carroll 639
Movements of radio-marked M. gallopavo silvestris increased our study
area to lands adjacent to Di-Lane Plantation WMA. The result was an overall
study-area size of 13,315 ha. Therefore, the WMA comprised 25% of the
overall study area. Although the WMA was managed extensively for wildlife,
it did mirror adjacent lands in many respects. Two large Zenaida
macroura L. (Mourning Dove) fields and smaller food plots corresponded to
agricultural areas, and Pinus spp. and hardwood stands were similar in
composition and structure to those of adjacent lands. Carya illinoinensis
(Wangenh.) K. Koch (pecan) orchards were also present on the WMA
similar to adjacent properties. In contrast to surrounding properties,
old-field complexes were more extensive and managed intensively, and
open-canopy planted Pinus spp. were more prevalent.
We delineated the study area into 7 habitat types (Table 1). Agricultural
areas were planted in Gossypium spp. (cotton), Glycine max (L.) Merr.
(soybeans), Arachis spp. (peanuts), or Zea mays L. (corn); however,
Gossypium spp. was the primary row crop. Pinus spp. plantations were
planted in P. taeda, generally 1793 trees/ha, and C. illinoinensis orchards
were treated as agricultural areas because of their intensive management.
Dominant hardwood species included Quercus nigra L. (water oak),
Quercus laurifolia Michx. (laurel oak), Quercus alba L. (white oak),
Quercus phellos L. (willow oak), Carya alba Nutt. ex Ell. (mockernut
hickory), Carya glabra Sweet (pignut hickory), Acer rubrum L. (red maple),
Nyssa sylvatica Marsh. (blackgum), and Liriodendron tulipifera L. (yellow
poplar). Mature Pinus spp. stands (> 20 years old) included either P. taeda or
Table 1. Habitat types delineated within the study area for analysis of habitat selection by radiomarked
Meleagris gallopavo silvestris, Burke County, GA, 1998 and 1999.
Habitat type % of areaA CompositionB
Hardwood 37 Stand composed of 50% hardwoods > 15 years old,
little herbaceous understory
Agriculture 22 Cultivated fields, soil-disturbed areasC, Carya
illinoinensis orchards, no or little herbaceous
understory other than crop
Closed-canopy planted Pinus spp. 16 Planted Pinus spp. stands at canopy closure, 8–20
years old, needle-laden ground cover
Field 13 Fallow areas, mowed areas, 1–4 year-old clearcuts,
prominent herbaceous understory
Old clearcut 6 Dense stand of mixed, natural regeneration 5–15
years old, stem densities significantly higher
Mature Pinus spp. 3 Stand composed of > 50% Pinus spp. > 20 years
old, little herbaceous understory
Open-canopy planted Pinus spp. 2 Thinned, planted Pinus spp. stands, < 5 year-old
planted Pinus spp., prominent herbaceous
AExcludes residential areas, paved roads, and open water deemed unsuitable for birds (1% of area).
BStands were 0.8 ha in size.
CIncludes clearcuts < 1 year-old that contain little vegetative ground cover and exposed soil.
640 Southeastern Naturalist Vol. 5, No. 4
Pinus echinata P. Mill. (shortleaf pine). They were characterized by little
herbaceous ground cover and generally juxtaposed near field areas.
We captured M. gallopavo silvestris with a rocket net in baited fields
(Wunz 1987) during January–March, 1998 and 1999. Each bird was fitted
with a uniquely-numbered leg band and a 110-g backpack-style radio transmitter
with a motion sensor (Telonics, Inc., Mesa, AZ). We sexed, weighed,
and aged each turkey before it was released from the trap site (Williams and
During each spring and summer, 3-element Yagi antennas and R4000
receivers (Advanced Telemetry Systems, Anoka, MN) were used by 2 observers
to record 2 bearings from permanent telemetry stations (n = 336) to each
radio-marked M. gallopavo silvestris (Cochran and Lord 1963). Greater than
90% of bearing sets were collected sequentially in 5 min in 1998; in 1999,
bearings were recorded simultaneously from stations that yielded the best
angle of intersection (a 90 degree angle being ideal). Bearing sets were
mapped in the field on 1:79-m scale aerial photos to ensure bearings crossed
and the estimated location was plausible (e.g., not in a pond or house).
Additionally, bearing sets were not recorded when bearings traversed multiple
habitat types. We avoided these circumstances by moving to transitions of
habitat types to ensure that the signal traversed vegetation of the same type.
Therefore, habitat-use error associated with the use of yagi antennas was
minimized, because we were able to identify the habitat block an individual
was located. Each M. gallopavo silvestris was located once in each of 3 time
periods (0600–1100 hr, 1101–1500 hr, and 1501–1800 hr) for a total of 3
locations/week. At least 6 locations/week were recorded for brooding hens for
2 weeks post-hatch. Bearing-error tests (White and Garrot 1990) were conducted
by hiding transmitters in known locations.
We digitized the study area in UTM coordinates on Digital Orthophoto
maps at a scale of 1:3000 m with ArcView software (Environmental
Systems Research Institute, Redlands, CA). Seven habitat types were delineated
by habitat blocks 0.8 ha in size. Permanent telemetry stations, visual
observations, and offset locations (birds within 100 m of observer) were
recorded with a GeoExplorer II hand-held global positioning system (GPS)
unit (Trimble Navigation Ltd., Sunnyvale, CA) in universal transverse
mercator (UTM) coordinates.
Meleagris gallopavo silvestris locations were determined using Location
of a Signal (LOAS) software (Ecological Software Solutions, Sacramento,
CA), and the program’s best biangulation estimator. Minimum convex polygon
home ranges were constructed for each M. gallopavo silvestris (mean =
33.5, SE = 1.4 locations) for summer ranges and 12 locations for brooding
ranges using ArcView software (Hooge and Eichenlaub 1997). We established
the summer season’s starting date by identifying the date on which 90%
of radio-marked hens began incubation. Summer home ranges for females
2006 J.J. Morgan, S.H. Schweitzer, and J.P. Carroll 641
only included post-incubation locations. The brooding period was a subset of
the summer season that included the first 2 weeks after hatching, because the
period is the most critical for recruitment (Vangilder 1992). Only hens
successfully hatching a nest were analyzed for the brooding period.
Summary statistics included mean, standard error of the mean (SE), and
sample size (N). We used compositional analysis to identify habitat preferences
by M. gallopavo silvestris within the study area and within their home
ranges (Aesbischer et al. 1993). Sample sizes were maximized for analyses
by pooling data across age classes and years (White and Garrot 1990).
Following Aebischer et al. (1993), proportions of habitat types within the
study area were compared to those within the minimum convex polygon, and
the proportion of radio locations in the habitat types were compared to the
proportions of habitat types within the minimum convex polygon. A multivariate
analysis of variance (MANOVA) was conducted to detect selection
of habitat types by M. gallopavo silvestris. Expected values were created by
randomization (Edgington 1980) for comparison with observed data. Nine
hundred ninety-nine iterations were run to generate a P-value (Carroll et al.
1995). If selection of habitat types was detected, then paired t-tests were
conducted to identify differences in use of each habitat type relative to other
habitat types. We used MANOVA to detect differences in habitat use between
sexes. Home-range sizes were compared by unequal variance t-tests
using MS Excel software (Microsoft Corporation, Redmond, WA). Significance
level for all statistical analyses was 0.05.
We collected 1715 locations (651 in 1998 and 1064 in 1999) from 28
male and 35 female M. gallopavo silvestris (Morgan 2000); however, 14
males and 22 females comprised the analysis because of mortality or
dropped/failed transmitters. Absolute mean bearing error was 8.7 ± 1.0
degrees (n = 158) in 1998 and 9.8 ± 0.7 degrees (n = 149) in 1999. Mean
bearing length was 439.2 ± 7.7 m (n = 3741). Seventeen percent of locations
were visual observations or offset locations. Tracking began in both years in
late March and ended 15 August 1998 and 31 July 1999. The onset of the
summer season was 11 May 1998 and 10 May 1999.
Summer home-range size for males (n = 14) averaged 320.6 ± 43.6 ha,
whereas that of females (n = 22) was 450.2 ± 62.4 ha. While brooding young,
home-range size of females averaged 101.6 ± 17.5 ha. Proportions of habitat
types available in the study area were different from the proportions of
habitat types within home ranges of males and females during the summer
(Table 2). Furthermore, the proportions of habitat types within the home
range were different than the proportions of radio-locations within the habitat
types for males and females (Table 2). Home ranges of brooding females
contained habitat types in different proportions than those in the study area
(L = 0.267, 4 df, P = 0.021), and proportions of radio-locations in habitat
types were different than the proportion of habitat types in the home range
642 Southeastern Naturalist Vol. 5, No. 4
(L = 0.112, 4 df, P = 0.013). Summer habitat use by males and females was
not different within the study area (L = 0.772, 6 df, P = 0.280) or within the
home range (L = 0.717, 6 df, P = 0.730).
During the summer, male M. gallopavo silvestris selected hardwood
habitats at the landscape (study area) and local (home range) scales. Comparisons
from the landscape to the local scale showed fields, closed-canopy
planted Pinus spp., and agriculture habitat types increasing in importance,
whereas the mature Pinus spp. habitat type declined in importance. The least
preferred habitat at both scales were old clearcuts and open-canopy planted
Pinus spp. stands (Table 2).
Females selected hardwood habitat types at both scales during the summer
as well. From the landscape to the local scale, field habitats decreased in
Table 3. Mean log-ratio difference matrix for all habitat pairings of proportions in minimum
convex polygons versus proportion of locations in each habitat for Meleagris gallopavo
silvestris hens (n = 10) during the brood period in Burke County, GA, 1999.
Hardwood Agriculture Field Pinus spp. Pinus spp.B
Habitat typeA (Mean, SE) (Mean, SE) (Mean, SE) (Mean, SE) (Mean, SE)
Hardwood – -5.22, 1.10 -0.98, 1.10 -0.64, 0.78 -3.74, 1.25
Agriculture +5.22, 1.10 – +5.84, 1.30 +3.58, 2.05 +2.70, 1.66
Field +0.98, 1.10 -5.84, 1.30 – +1.15, 1.49 -3.13, 1.29
Mature Pinus spp. +0.64, 0.78 -3.58, 2.05 -1.15, 1.49 – -2.94, 2.08
Planted Pinus spp. +3.74, 1.25 -2.70, 1.66 +3.13, 1.29 +2.94, 2.08 –
RankC 1 5 3 2 4
AA negative log-ratio difference value indicates that the relative use of the row habitat was more
than the column habitat. A positive value indicates the opposite.
BOpen- and closed-planted pine habitat types were combined for the analysis because of small
CRanks determined by comparing the relative use of each habitat against all other habitats.
Smallest ranking indicates the most used habitat relative to abundance.
Table 2. Compositional analysis of Meleagris gallopavo silvestris habitat selection (1 = most
relative use) within the study area and home range, Burke County, GA, 1998 and 1999.
Males (n = 14) Females (n = 22)
Habitat type Study area Home range Study area Home rangeA
Agriculture 5 4 5 5
Closed-canopy planted Pinus spp. 4 3 4 2
Field 3 2 2 4
Hardwood 1 1 1 1
Mature pine 2 5 3 3
Old clearcut 6 6 6 5
Open-canopy planted Pinus spp. 7 7 7 5
Wilk’s Lambda 0.067 0.024 0.103 0.036
P-valueA 0.006 < 0.001 < 0.001 < 0.001
AAgriculture, old clearcut, and open-canopy planted Pinus spp. all tied for the least relative use
in the females' home ranges.
BIf P 0.05, then habitat use was not random.
2006 J.J. Morgan, S.H. Schweitzer, and J.P. Carroll 643
importance and closed-canopy planted Pinus spp. increased in importance
(Table 2). Least important habitat types to females at both scales were
agriculture, old clearcut, and open-canopy planted Pinus spp. Females with
poults used hardwoods most frequently and agricultural areas least frequently
at the local scale (Table 3).
Meleagris gallopavo silvestris use of Pinus spp. plantations in the Southeast
has been widely studied and results vary. The preponderance of research
has focused on pine-dominated areas. Habitat use changed by season, sex,
and plantation age and stem density. Spring and summer use varied the most
among studies making comparisons difficult. For example, young plantations
(< 4 years old) were not preferred habitat in any season, but were used
occasionally in proportion to availability in Arkansas (Wigley et al. 1985).
Conversely, Bidwell et al. (1989) found M. gallopavo silvestris use of
plantations limited to the young age class (1–4 years old) in Oklahoma,
because vegetation composition was similar to small openings. Middle-aged
(4–14 years old) stands generally were used by males in the summer (Exum
et al. 1987, Wigley et al. 1985), whereas older stands (> 14 years) were
selected by hens in the spring (Exum et al. 1987). In the summer, males and
females used and selected closed-canopy planted Pinus spp. habitats in our
study, despite having well-distributed field and hardwood habitats.
The differences between studies may be based on cover and food requirements.
Pinus spp. habitats at canopy closure provide overhead cover and
open understories that may afford M. gallopavo silvestris protection from
weather and predators. Juxtaposition may also be a factor (Phalen et al.
1986). Closed- and open-canopy Pinus spp. are often bordered by firebreaks,
so M. gallopavo silvestris have the benefit of cover and feeding areas in
close proximity. Many Pinus spp. plantations on our study area were adjacent
to fields and hardwoods offering travel ways to feeding and roosting
sites. Finally, the landscape context of our study was unique compared to
previous research. In pine-dominated study areas, M. gallopavo silvestris
have little opportunity to utilize a variety of habitats (i.e., Pinus spp. plantations
and non-pine habitats).
M. gallopavo silvestris summer home ranges in our study area were
generally smaller than those reported by other studies (Badyaev et al. 1996a,
Bidwell et al. 1989, Everett et al. 1980, Hurst et al. 1991, Smith et al. 1988).
Home-range size may be related to habitat quality (Exum et al. 1987, Porter
1977). Historic definitions summarized by Porter (1992) commonly identified
ideal habitat as woodland areas consisting of 5 to10% grassy openings. Our
study area was interspersed with fields totaling 13% of the available habitat.
Badyaev et al. (1996b) suggested that home ranges of females during spring
were large because of extensive movements in search of nesting sites.
Hillestad and Speake (1970) identified the availability of pastures as a factor
affecting small range size in brooding hens. Home-range sizes of brooding
644 Southeastern Naturalist Vol. 5, No. 4
hens in our study were similar to those of other studies (Burk et al. 1990,
Phalen et al. 1986). An abundance of nesting and brooding habitat on our
study area may have resulted in small female home ranges. Furthermore,
home-range sizes are smaller when a diversity of forest habitats and openings
are available for food and cover (Korschgen 1967, Speake et al. 1975), and our
study area typified these conditions.
Relative to all available habitat types, mature hardwood stands were the
most frequently used habitat in the summer. Their importance to M.
gallopavo silvestris has been documented repeatedly. Hardwoods are selected
in fall and winter (Holbrook et al. 1987, Palmer et al.1993) providing
forage and cover (Kennamer et al. 1980, Porter 1992). Strips of hardwoods
furnish travel corridors between feeding areas and serve as roosting and
loafing sites (Gehrken 1975, Holbrook et al. 1985). In spring and summer,
hardwood stands were selected consistently and supplied M. gallopavo
silvestris with hard and soft mast, forbs and grasses, and insects (Miller et al.
1999). Hardwood stands in our study area contained little midstory or
shrubby vegetation, and our findings further support the value of similar
hardwood stands to M. gallopavo silvestris populations.
M. gallopavo silvestris generally avoided agricultural fields and old
clearcuts as expected. Kurzjeski and Lewis (1990) documented low use of
agricultural areas in Missouri during the spring and summer. Intensive
agriculture for Gossypium spp. and other row crops included the application
of herbicides and insecticides. Row crops were nearly weedless and likely
supported few invertebrates making them unsuitable feeding areas for M.
gallopavo silvestris. Old clearcuts were characterized by dense vegetation
that would impede the movements of M. gallopavo silvestris and were too
young to produce hard mast (Wunz and Pack 1992).
Management objectives for M. gallopavo silvestris often focus on habitats
promoting reproduction, and openings (i.e., fields and pastures)
provide the highest quality brood-rearing habitat for M. gallopavo
silvestris (Healy and Nenno 1983, Lewis 1964, Peoples et al. 1995, Speake
et al. 1975). On our study area, we found that brooding hens showed the
strongest selection for hardwoods. Phalen et al. (1986) reported that most
broods moved to bottomland hardwoods in Mississippi, but field habitats
were only available on the periphery of the pine-dominated study area.
Conversely, hardwoods selected by brooding hens in our study were generally
in uplands, but were adjacent to fields. Everett et al. (1985) reported
that males and females in Alabama moved away from bottomland areas to
open habitats that provided grasses for feeding and brood rearing in the
summer, and our results support their conclusion.
Brooding hens’ selection for fields and mature Pinus spp. further supported
the value of upland areas in our study area. Use of upland planted
Pinus spp. habitats by brooding hens was low, contrary to findings in pinedominated
habitats in Mississippi (Burke et al. 1990). In coastal plain Pinus
spp. forests of Georgia and Florida, hens selecting openings (e.g., fields)
2006 J.J. Morgan, S.H. Schweitzer, and J.P. Carroll 645
were more successful at raising poults (Peoples et al. 1995). Upland habitats
in our study contained fields, but hens were most often located within
Intuitively, the USDA’s recommendations for thinning Pinus spp. stands
for CRP re-enrollment should promote use by hens in the summer. Thinning
increases forage production and vertical structure in planted Pinus spp.
stands (Conroy et al. 1982, Lewis 1999, Melchiors 1991) and should increase
the value of the habitat for brood rearing. Our research does not
support that premise, but our study area was characterized by only 2% of
open-canopy planted Pinus spp. and had well-interspersed fields.
We found selection of closed-canopy Pinus spp. stands by males and
females in the summer when other habitat types (i.e., fields and hardwoods)
were readily available. Brooding hens used planted Pinus spp.
more than agricultural fields, although neither were highly selected. Our
study area was characterized by Pinus spp. plantations in an agricultural
context that was more representative of the landscape influenced by CRP
Pinus spp. plantings than previous research. The area included interspersed
fields and large hardwood drains adjacent to Pinus spp. plantations. Therefore,
foraging and brood-rearing habitat may not have been limiting. Pinus
spp. plantations may have afforded M. gallopavo silvestris loafing sites
and served as travel corridors to feeding and roosting areas. Meleagris
gallopavo silvestris are highly adaptable and mobile, and the replacement
of agricultural fields with planted Pinus spp. stands may have improved
habitat quality when fields and mature hardwood drains are available.
Further research on brood rearing in mixed agricultural and silvicultural
landscapes, particularly focused on planted Pinus spp. management typical
of CRP (e.g., thinning requirements), may further identify the effect of
CRP Pinus spp. on M. gallopavo silvestris.
Funding for this research was provided by the National Wild Turkey Federation
through funding from an anonymous donor to The National Fish and Wildlife
Foundation. Additional support was provided by the Game Management Section of
the Georgia Department of Natural Resources’ Wildlife Resources Division and by
the University of Georgia’s Warnell School of Forestry and Natural Resources.
Special thanks to F. Schroeder and K. Seginak for their tireless field work and C.G.
White for statistical assistance.
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