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Southeastern Naturalist
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2018 Vol. 17, No. 3
SOUTHEASTERN NATURALIST
2018 17(3):371–380
Eastern Wild Turkey Roost-site Selection in a
Fire-maintained Longleaf Pine Ecosystem
Indrani Sasmal1,*, Eric L. Kilburg1, Christopher S. DePerno1, M. Colter Chitwood2,
Marcus A. Lashley3, Bret A. Collier4, and Christopher E. Moorman1
Abstract - Night-time roosting in Meleagris gallopavo (Wild Turkey) is a quotidian activ-
ity that minimizes vulnerability to predators and weather. Roost-site selection in managed
Pinus palustris (Longleaf Pine) communities is poorly documented. We assessed roost-site
selection by comparing use and availability of vegetation types at the individual female
Wild Turkey home-range level. We monitored 14 Wild Turkeys from February 2011 to June
2012. The Wild Turkeys did not use vegetation types within the estimated home ranges
for roosting in proportion to availability (χ² = 601.696, P < 0.001). Female Wild Turkeys
roosted in the upland Longleaf Pine in proportion to availability, selected for lowland hard-
wood, and avoided upland hardwood patches. We documented that roost-site availability is
not likely a limiting factor in managed Longleaf Pine forests.
Introduction
Roosting locations for Meleagris gallopavo silvestris Vieillot (Eastern Wild
Turkey; hereafter, Wild Turkey) limit vulnerability to predation and can provide
refugia from poor weather conditions (Byrne et al. 2016, Kilpatrick et al. 1988,
Ludwig 2012, Porter 1978). Hence, roosting sites are a critical habitat component
for Wild Turkeys (Bailey and Rinell 1967, Chamberlain et al. 2000). The structure
and composition of Wild Turkey roosting locations are similar across the species’
range (Kimmel and Zwank 1985, Still and Baumann 1989, Zwank et al. 1988). In
the southeastern US, Wild Turkey roost sites often are in lowland hardwood stands
adjacent to permanent water or in Pinus (pine)–hardwood stands (Chamberlain et
al. 2000, Kimmel and Zwank 1985, Miller et al. 1999, Zwank et al. 1988).
Although Wild Turkey roost-site selection has been documented in a variety
of community types (Chamberlain et al. 2000, Kilpatrick et al. 1988, Tzilkowski
1971), information on roost-site selection in frequently burned Pinus palustris Mill.
(Longleaf Pine) communities is lacking. Longleaf Pine communities represent one
of the most diverse ecosystems in the temperate zone and commonly are restored
and maintained with frequent, low-intensity prescribed re (Drew et al. 1998, Fill
et al. 2012, Lashley et al. 2015). However, homogeneous application of burning
1Fisheries, Wildlife, and Conservation Biology Program, Department of Forestry and
Environmental Resources, North Carolina State University, Raleigh, NC 27695. 2Wild-
life Biology Program, Department of Ecosystem and Conservation Sciences, University
of Montana, Missoula, MT 59812. 3Department of Wildlife, Fisheries, and Aquaculture,
Mississippi State University, Mississippi State, MS 39762. 4School of Renewable Natural
Resources, Louisiana State University, Baton Rouge, LA 70803. *Corresponding author -
bulirchithi@gmail.com.
Manuscript Editor: Robert Carter
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techniques, return interval, and season of burn can decrease compositional and
structural heterogeneity of plant communities in frequently burned Longleaf Pine
communities by differentially promoting the prevailing vegetation type (Longleaf
Pine woodland) and suppressing less-prominent hardwood inclusions (Lashley et
al. 2014). Thus, if mature upland hardwoods provide the best roosting cover, cur-
rent prescribed re regimes may be problematic for Wild Turkeys.
Historically, much of the southeastern US burned frequently, and experimentation
with prescribed re has produced vegetation conditions that benet Wild Turkeys by
providing more diverse or more abundant food and higher-quality nesting cover (Cox
and Widener 2008, Kilburg et al. 2015, Knapp et al. 2009, Lashley et al. 2015). Yet,
little is known about Wild Turkey roost-site selection in frequently burned Longleaf
Pine forests—a landscape where lowland hardwood availability often is limited.
Therefore, our objective was to assess roost-site selection by female Wild Turkeys in
frequently burned Longleaf Pine woodlands in central North Carolina.
Field-Site Description
We evaluated female Wild Turkey roost-site selection at Fort Bragg Military
Installation (hereafter, Fort Bragg) in the Sandhills physiographic region of North
Carolina. The Sandhills region is characterized by variably deep, well-drained, and
sandy soils, with xeric uplands and hillside seeps that feed numerous blackwater
streams (Sorrie et al. 2006). Frequent re and variable soil moisture produced
several vegetation types at Fort Bragg (Sorrie et al. 2006), including lowland
hardwood (10% of the land area), upland hardwood (4%), upland pine (69%), and
non-forested (17%) (Lashley et al. 2014).
Lowland hardwoods contained Acer rubrum L. (Red Maple), Liquidambar
styraciua L. (Sweetgum), Liriodendron tulipifera L. (Tulip-poplar), and Nyssa
sylvatica Marsh. (Blackgum), forming generally closed canopy stands along per-
manently owing streams. Dense thickets of Ilex spp. (gallberries), Lyonia spp.
(Fetterbush), and Smilax spp. (greenbriers) comprised the understory. Xeric hard-
wood species (primarily Quercus spp. [oaks]) dominated upland hardwood areas. A
Longleaf Pine overstory, with an understory of Aristida stricta Michx. (Wiregrass),
Gaylussacia dumosa (Andrews) Torr. & A. Gray (Dwarf Huckleberry), Q. laevis
Walter (Turkey Oak), and Q. marilandica Münchh. (Blackjack Oak), dominated the
uplands. Upland pine stands were burned every 3 years during the growing-season
(i.e., April–August) to control woody-stem encroachment to the forest midstory
in accordance with management objectives for the endangered Leuconotopicus
borealis (Vieillot) (Red-cockaded Woodpecker). Non-forested vegetation oc-
curred primarily in areas with military activity (hereafter, military-activity zones),
including artillery-ring points, aerial-drop zones, and artillery-impact areas. Mili-
tary-activity zones were all sparsely vegetated and dominated by grasses and forbs,
including non-native Eragrostis curvula (Schrad.) Nees (Weeping Lovegrass) and
Lespedeza cuneata (Dum. Cours.) G. Don (Sericea Lespedeza). Drop zones were
burned and mowed annually or biennially to reduce woody vegetation for the safety
of paratroopers; these areas provided no roosting cover for Wild Turkeys.
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Methods
We captured Wild Turkeys by rocket net during February–April 2011 and
January–March 2012. In 2011, we fitted each captured female Wild Turkey with
an 85-g micro GPS-data logger (Model G1H271; Sirtrack LTD, Havelock North,
New Zealand) programmed to obtain 4 locations daily (every 6 h, beginning at
00:00:00). We set the fix rate to optimize relocation frequency with data-logger
battery life to ensure the devices could collect data for >1 year. Data loggers were
equipped with radio transmitters and programmed to store relocation coordinates
onboard (Guthrie et al. 2011). All capture and handling protocols were approved
by North Carolina State University Institutional Animal Care and Use Committee
(#10-149-A).
Kernel methods may perform poorly with large data sets when using common
methods of determining the smoothing parameter (h) (Getz and Wilmers 2004,
Hemson et al. 2005). Hence, we used dynamic Brownian-bridge movement models
(Kranstauber et al. 2012) to estimate year-round utilization distributions (UDs) for
female Wild Turkeys. We based UDs on the movement tracks of each individual
(February 2011–June 2012) using R package move (Kranstauber and Smolla 2016)
in Program R version 3.4.2 (R Core Team 2016). The dynamic Brownian-bridge
movement model incorporates the behavioral heterogeneity of the movement pro-
cess (Horne et al. 2007, Kranstauber et al. 2012) and quanties individual-space
use using individual behavioral information. We used a GPS-error estimate of 20
(Byrne et al. 2014, Guthrie et al. 2011), a raster value of 100, and time-step value of
60 (equivalent to 1 hour) with a moving-window size of 29 relocations (equivalent
to 7 d) with a margin of 9 relocations over full tracks of each Wild Turkey.
We assessed roost-site selection within female Wild Turkey ranges using re-
source-selection Design III (Manly et al. 2002). We included only 1 nocturnal relo-
cation per female to ensure that we quantied only a single roost-location per night.
Using ArcGIS 10.3.1 (ESRI, Redlands, CA) and Fort Bragg’s vegetation-type layer
(Fig. 1), we determined percentages of each forested vegetation type (lowland
hardwood, upland pine, and upland hardwood) within the estimated home-range
of each individual Wild Turkey. We assessed roost-site selection by comparing use
and availability of vegetation types within each estimated home-range (Manly et
al. 2002). We dened use as the number of roost locations in a particular vegetation
type, and availability as the percentage of that vegetation type available within the
individual range. We calculated selection ratios and chi-square values to estimate
the overall deviation from random use using program R version 3.2.4 (R Core Team
2016) and the 'adehabitat' package (Calenge 2006). Selection ratios (ŵ) indicated
selection if estimates differed from 1, and we computed ratios for each vegetation
type and individual as the ratio of used proportion to available proportion (Calenge
and Dufour 2006). Selection for vegetation types was indicated if the lower limit of
the 90% condence interval (CI) of ŵ was >1, whereas selection against vegetation
types was indicated if the upper limit of the 90% CI of ŵ was <1. Use in proportion
to availability (neutral selection) was indicated if the 90% CI of ŵ contained the
value 1 (Manly et al. 2002).
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We generated a minimum convex polygon (MCP) area of all roost locations
using Home Range Tool version 2 in ArcGIS. We generated equal numbers of ran-
dom locations within the buffered MCP area of all roost locations, which we used
to delineate the boundaries for vegetation-type analysis. We measured distances
from roosts and random locations to rebreaks/roads, streams, and military-activity
zones using the proximity tool in ArcGIS 10.3.1. We used paired t-tests to assess
whether distance from rebreaks/roads, streams, and military-activity zones dif-
fered between Wild Turkey roost sites and random locations at the 90% level of
signicance (α = 0.1).
Results
We recovered data from 14 GPS tagged Wild Turkeys (13 in 2011 and 1 in 2012),
which recorded 11,655 relocations (mean = 833) between February 2011 and June
2012. Average annual home-range size was 8.54 km2 (SE = 62; Table 1). We recorded
2610 roost locations; not all vegetation types within the 95% home-range estimates
were used in proportion to availability (χ² = 601.696, P < 0.001; Table 2). Wild Tur-
keys used the upland pine (90% CI = 0.68–1.03) in proportion to availability, whereas
Figure 1. Map of forest types used to study female Wild Turkey roost-site selection from
February 2011 to June 2012 at Fort Bragg Military Installation, NC. The white areas within
the forest-type map represent non-forested areas.
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they avoided upland hardwoods (90% CI = 0.44–0.72) and selected for lowland hard-
woods (90% CI = 2.45–4.3) (Table 3). Random locations were an average of 245.75
m (SE = 3.78) from a stream, 96.24 m (SE = 2.64) from a rebreak/road, and 357.29
m (SE = 8.33) from a military-activity zone. Female Wild Turkey roost locations
were an average of 238.21 m (SE = 3.76) from a stream, 112.9 m (SE = 3.75) from a
rebreak/road, and 490.32 m (SE = 10.95) from a military-activity zone. Compared
to random locations, female Wild Turkeys selected roost sites farther from rebreaks/
roads (P < 0.001) and military-activity zones (P < 0.001). Distance to streams was
similar (P = 0.16) between roost sites and random locations.
Table 1. Female Wild Turkey (n = 14) home-range (95%) size estimated using a dynamic Brownian-
bridge movement model, Fort Bragg Military Installation, NC, February 2011–June 2012.
Turkey ID 95% home-range (km2)
851 11.644
800 11.629
701 8.267
650 12.152
551 5.911
450 6.295
371 5.351
350 6.452
311 9.213
251 6.407
171 8.498
123 7.851
91 10.787
21 9.149
Table 3. Roost-site selection, including selection ratios (ŵ), standard errors (SE), and 90% condence
intervals (CI), by vegetation type for female Wild Turkeys (n = 14) at Fort Bragg Military Installation,
NC, February 2011–June 2012. Selection for roosting in a vegetation type is indicated by a CI above
1, selection against by a CI below 1, and use in proportion to availability (i.e., neutral selection) by
a CI overlapping 1.
Vegetation type ŵ SE Lower CI Upper CI
Upland pine 0.85 0.08 0.68 1.03
Upland hardwood 0.58 0.07 0.44 0.72
Lowland hardwood 3.37 0.43 2.45 4.30
Table 2. Available (%) and used (%) forested vegetation types for roosting female Wild Turkeys (n =
14) at Fort Bragg Military Installation, NC, February 2011–June 2012.
Vegetation type* Available (%) Used (%)
Upland pine 21.69 25.14
Upland hardwood 13.36 9.68
Lowland hardwood 3.12 16.21
*Remainder was not forested (i.e., military-drop zones, ring zones, water bodies, and roads)
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Discussion
Female Wild Turkeys frequently roosted in upland Longleaf Pine, but the con-
dence interval for the selection ratio only slightly overlapped zero, indicating
some avoidance of the vegetation type. Animals adjust their space use according
to resource availability to optimally exploit the resources (Fauchald 1999, Turchin
1991), and it has been demonstrated that roosting sites are selected according to
availability of potential sites on the landscape (Byrne et al. 2015). Thus, the sheer
abundance of Longleaf Pine woodland available on the Fort Bragg landscape could
explain its value as potential roosting cover for Wild Turkeys. Finer-scale location
and vegetation data are needed to help determine the degree of selection for roosts
within upland pine stands. For example, Chamberlain et al. (2000) noted 2 possible
scenarios for roost-site selection that could be relevant for explaining Wild Turkey
roosting behavior in upland Longleaf Pine stands at Fort Bragg: (1) females forag-
ing in upland pine stands may simply y up to roost in the nearest roosting cover at
the end of the day, or (2) females may be using their daily movements through the
upland pine stands to arrive at predetermined roosting sites by evening. Regard-
less of these 2 scenarios, Wild Turkeys likely use upland pine for roosting because
of sparse understory (Palmer et al. 1996) and the potential protection provided by
conifers against harsh weather (Bailey and Rinell 1967, Kilpatrick et al. 1988).
Similar to elsewhere in the species’ range, female Wild Turkeys in our study
selectively roosted in lowland hardwood habitat, but individuals avoided upland
hardwood patches. Lowland hardwood covered ~10% of the land area on Fort
Bragg but offered critically important roosting cover. Although we did not examine
roost-site selection at the level of individually selected trees, Wild Turkey use of
lowland hardwoods for roosting likely suggests selection for taller hardwood trees.
Lowland hardwood areas generally contained tall non-oak hardwoods (e.g., Black-
gum, Tulip-poplar) and interspersed P. taeda L. (Loblolly Pine) (Prince et al. 2016).
At Fort Bragg, mature hardwoods were removed mechanically in some upland
stands, potentially restricting the tallest hardwoods to riparian areas that did not
burn or along rebreaks that provided a re shadow (Lashley et al. 2014). Indeed,
most of the largest-diameter hardwoods and pines were located in riparian areas
at Fort Bragg, and those areas generally contained low densities of reproductively
mature oak trees (Lashley et al. 2014). Given the potential importance of large
hardwoods for Wild Turkey roosting coupled with the known importance of hard
mast to the Wild Turkey diet (Dickson 2001), management actions in Longleaf
Pine communities (i.e., frequent res, chemical or mechanical treatments) that limit
the abundance and distribution of mature hardwoods could negatively affect Wild
Turkey roost availability, especially if the re regime is applied in a way that limits
re shadows that promote succession of hardwoods to maturity in this ecosystem
(Lashley et al. 2014).
Roosting Wild Turkeys avoided the military-activity zones and rebreaks/
roads, but demonstrated no selection for or avoidance of water sources. The birds
likely avoided military-activity zones because they generally lacked trees suitable
for roosting, except along the edges of the openings. However, it is also probable
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that frequent anthropogenic disturbance in the military-activity zones and along
rebreaks/roads may have deterred roosting Wild Turkeys. Although some stud-
ies have suggested that Wild Turkeys roost near water sources (Boeker and Scott
1969, Chamberlain et al. 2000, Kilpatrick et al. 1988), we did not detect selection
or avoidance for roosting near or over water. Selection of roost sites in proximity
to water in other studies might be an artifact of the improved foraging resources
nearby (Crockett 1973), which might explain why we did not detect a similar effect.
Moreover, data limitations precluded our ability to test for seasonal trends in roost-
site selection, which could have reduced our ability to identify seasonal importance
of water sources.
Movements and behavior of Wild Turkeys can be inuenced by roost sites (Gross
et al. 2015), so maintenance of roost-site availability in Longleaf Pine communi-
ties is warranted. At Fort Bragg, limited availability of tall hardwoods coupled with
anthropogenic disturbance appeared to have the combined effect of limiting the
areas that Wild Turkeys selected for viable roost sites. Although frequent re has
numerous ecological benets for Wild Turkeys and other taxa in the Longleaf Pine
ecosystem, managers will need to consider the compounding effects of additional
management actions (e.g., mechanical removal of hardwoods) that could further re-
duce the hardwood component on the landscape. For example, Streich et al. (2015)
determined that frequent re (≤2-y return-interval) was compatible with conser-
vation of Wild Turkey nest-site and brood ground-roost cover but that managers
should carefully consider removal of hardwoods, particularly in riparian areas, due
to their importance to hens and broods. Also, frequent re can eliminate species that
produce eshy fruits, which are an important food resource for Wild Turkeys (Lash-
ley et al. 2015, 2017). Moreover, removal of relic mature oaks within forest stands
may be particularly problematic without adjusting concurrent re regimes to allow
oak succession in re shadows (Lashley et al. 2014). Thus, managers of the Long-
leaf Pine ecosystem should promote heterogeneous landscape conditions includ-
ing re-maintained uplands as well as lowland hardwoods that are less frequently
burned to provide roosting and nesting cover for Wild Turkeys, while simultaneous-
ly allowing the restoration and maintenance of habitat conditions for other wildlife
species associated with the ecosystem (Kilburg et al. 2014, 2015; Prince et al. 2016,
Sasmal et al. 2017). To conserve Wild Turkey roosting cover in the Longleaf Pine
ecosystem, resource managers should strive to protect lowland hardwoods and cre-
ate a mosaic of upland hardwoods that include both recently burned as well as less
recently burned sections, allowing for the regeneration and maturation of oaks and
other hardwoods in areas of low topography and mesic areas near streams (i.e., in
re shadows; Prince et al. 2016).
Acknowledgments
Funding for this research was provided by the US Department of Defense. We thank
M. Broadway, M. Nunnery, B. Peterson, and B. Sherrill for eld assistance. We thank A.
Shultz and J. Jones of the Fort Bragg Wildlife Branch for providing trapping equipment and
eld assistance. The North Carolina Wildlife Resources Commission and US Department of
Agriculture Wildlife Services assisted with trapping and provided equipment.
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2018 Vol. 17, No. 3
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