Forest Ecology and Management 500 (2021) 119646 Available online 27 August 2021 0378-1127/Published by Elsevier B.V. Alter fire timing to recouple forage nutrients with herbivore nutrient demands Rainer A. Nichols a , * , Steve Demarais a , Bronson K. Strickland a , Marcus A. Lashley a , b a Department of Wildlife and Fisheries, Mississippi State University, P.O. Box 9690, MS, USA b Department of Wildlife Ecology and Conservation, University of Florida, 110 Newins Ziegler Hall, Gainesville, FL 32611, USA A R T I C L E I N F O Keywords: Dormant season Fire timing Forage quality Forage quantity Growing season Nutritional carrying capacity A B S T R A C T Forests typically have limited high quality nutrition available to meet the seasonal needs of white-tailed deer ( Odocoileus virginianus ) during summer. Prescribed fire is used to enhance deer habitat in pyric-disclimax com- munities but is normally applied during the dormant season. Historically, lightning-ignited fires in pine systems occurred during summer in the southeastern US, but fire ignited by humans occurred at all times of the year and was most common in fall/winter. The different timing of prescribed fire may be important when trying to couple forage resources with nutritionally demanding periods for herbivores. We evaluated fire-timing effects (dormant season, March; growing season, June; and unburned control) on nutrient availability (crude protein, % phos- phorus, and biomass) and nutritional carrying capacity (NCC) for moderate-high use forages of white-tailed deer during summer of 2018 and 2019 in nine mid-rotation loblolly pine stands. Fire increased nutrient content the year of the burn and the increase persisted into the second growing season following growing season fire. Dormant season fire increased protein- and phosphorus- based NCC for two growing seasons but growing season fire reduced biomass during the first year and delayed NCC improvement until the second year. Prescribed fire can be timed to manipulate forage quality, quantity, and improve NCC to recouple forage nutrient supply with herbivore seasonal nutrient demands. We recommend land managers consider diversifying fire timing across the landscape in pyric disclimax systems to support the changing nutrient requirements of large herbivores. 1. Introduction Historically, fire played a critical role in many plant communities (Bond and Keeley, 2005) based on lightning strikes and anthropogenic ignition by Native Americans (Outcalt, 2008). Widespread fire sup- pression (Fowler and Konopik, 2007) altered ecological processes (Brockway et al., 2009), and pyric disclimax ecosystems shifted to alternative stable states (Brennan et al., 1998, Fowler and Konopik, 2007). Current anthropogenic fires set during the dormant season are phenologically mismatched to that of lightning fires (Knapp et al., 2009). Prescribed fire is considered a cost-effective management tool for land managers who seek to enhance white-tailed deer ( Odocoileus vir- ginianus ) habitat. It promotes and maintains plant community compo- sition that also provides foraging patches for many other wildlife species (Lashley et al., 2015a, Harper et al., 2016, Sittler et al., 2019) by reducing woody dominance and increasing forb and grass cover in the understory (Brennan et al., 1998, Iglay et al., 2010). It also ephemerally increases quality and quantity of forage for terrestrial herbivores by improving germination conditions and stimulating nutritious re-growth (Lewis et al., 1982, Dyke and Darragh, 2007, Lashley et al., 2011). However, gains in forage quality and quantity are influenced by when fire is implemented. Fire regimes in the Southeast are commonly implemented in the dormant season (Knapp et al., 2009), so varying the timing of prescribed fire could be an important consideration to improve species richness, diversity, and abundance of plants (Jeffrey et al., 1998, Lashley et al., 2015b). Fire during the growing season stimulates forage quality and decreases forage quantity during the same growing season (McCord et al., 2014, Lashley et al., 2015a, Sittler et al., 2019). Dormant season fire tends to increase forage production during the first growing season but does not increase quality because ephemeral gains of nutrition are typically lost within 3 – 4 months following fire (Lewis et al., 1982, Eby et al., 2014). Nutritional values of many forages in the southeastern U.S. typically decrease during the late summer (Jones et al., 2008) in contrast to * Corresponding author at: 166 Notting Hill Pl., Canton, MS 39046, USA. E-mail address: rainer.nichols.nfg@mail.mil (R.A. Nichols). Contents lists available at ScienceDirect Forest Ecology and Management journal homepage: www.elsevier.com/locate/foreco https://doi.org/10.1016/j.foreco.2021.119646 Received 1 June 2021; Received in revised form 17 August 2021; Accepted 20 August 2021 Forest Ecology and Management 500 (2021) 119646 2 increasing nutrient demands of deer during this time (Fig. 1). By the time many plants mature, the majority of antler growth has completed. However, these increased nutritional demands still occur for gestating and lactating does. This seasonal mismatch between nutrient availabil- ity and animal needs in the Southeast may result in a summer nutritional stress period. Dormant season prescribed fire is commonly used to in- crease deer nutritional carrying capacity (NCC) (Edwards et al., 2004, Iglay et al., 2006, Iglay et al., 2010, Lashley et al., 2011), but knowledge of growing season fire effects on summer NCC is lacking (Nanney et al., 2018). We used the largest herbivore on the southeastern US landscape (white-tailed deer) as a model to demonstrate how fire timing might influence nutrient availability. Prescribed fire regimes used to benefit deer also benefit other wildlife species, such as red-cockaded wood- pecker ( Leuconotopicus borealis ) and gopher tortoise ( Gopherus poly- phemus ), that rely on early successional plant communities within pine systems (Waller and Alverson, 1997). So, managing forests with stra- tegic timing of fire may create a more diverse landscape management by reducing hardwood encroachment and increasing heterogeneity of plant species and structure. To better understand the effects of fire timing on nutrient avail- ability, we compared growing season forage quality, quantity, and NCC for white-tailed deer during two growing seasons following prescribed fires set during dormant season (March), growing season (June), and un- burned controls. Given that fire produces ephemeral gains in forage quality (Lewis et al., 1982, Eby et al., 2014), we predicted growing season fire will increase forage quality during the first growing season and nutritional gains from both fire treatments be lost in the second growing season. Given that biomass production is dependent on when fire is implemented (McCord et al., 2014, Lashley et al., 2015a, Sittler et al., 2019), we predicted dormant season fire will produce more biomass during the first growing season than growing season fire and both fire treatments will increase quantity of biomass during the second growing season. 2. Methods 2.1. Study area We conducted this study in nine mid-rotation (~21 years old), thinned loblolly pine ( Pinus taeda ) stands in Monroe and Oktibbeha counties, Mississippi, USA (Fig. 2). The pine stands averaged 12.3 ha with a pine basal area of 22 m 2 ha 1 , ranging from 16-27 m 2 ha 1 Prescribed fire had not been conducted in these stands for at least 5 years. Dominant soil types on study areas in Oktibbeha County were Wilcox silty clay loam (Alfisols), Falkner and Longview silt loam (Alfi- sols), Urbo silty clay (Entisols), and Prentiss silt loam (Ultisols). Domi- nant soil types were Vaiden silty clay (Alfisols) in Monroe County. Rainfall during the entirety of this study was ~309 cm (U.S. Climate Data). 2.2. Treatment design Each of the nine pine stands (blocks) contained the randomly assigned treatments: dormant season fire, growing season fire, and un- burned control. Experimental units within blocks ranged from 2.5 to 8.0 ha and were separated by plowed fire breaks. Treatment units were approximately the same size within each block. We conducted dormant season fires during March 2018 (March 8 – 15) and growing-season fires during June 2018 (June 4 – 27). We conducted prescribed fire under weather conditions approved within a Mississippi burn permit, with relative humidity averaging 28% (range 25 – 32%) during March and 46% (range 34 – 59%) during June and wind speeds varying from 8 to 24 kph. Pyrometers were placed systematically across each experimental unit to measure fire intensity. Fire intensity averaged 227 ◦ C for dormant season fires and 272 ◦ C for growing season fires. We estimated biomass of moderate-high use deer forages (Warren and Hurst, 1981, Miller and Miller, 1999, Lashley et al., 2011) during August-September of 2018 and 2019 using 10, 1 m 2 enclosure cages situated at 10 systematically random locations across each experimental unit. We collected all leaves and growing tips of forages to represent consumable biomass following guidelines of Lashley et al. (2014). We dried samples in a convection oven at 47 ◦ C until constant mass was reached and sent them to Custom Laboratory (3068 State Hwy 37, Monett, MO 65708), a National Forage Testing Association certified lab, for a wet chemistry nutritional analysis of crude protein (CP) and phosphorus (P). We calculated treatment-specific, growing season, nutritional car- rying capacity for a mean-mixed diet quality of 14% CP (Jones et al., 2009, Iglay et al., 2010, Lashley et al., 2011) and 0.25% P (McEwen et al., 1957, Lashley et al., 2015c) to represent minimum nutrient re- quirements for a lactating doe with one fawn using the explicit nutri- tional constraints model (Hobbs and Swift, 1985). We assumed deer consumed 1.36 kg of dry-matter forage per day (French et al., 1956, Fig. 1. Declining plant quality with increasing nutritional needs from spring to summer in the Southeast. This trend signifies the late summer stress nutritional period that white-tailed deer undergo. Fig. 2. Location of nine pine stands on three properties in Monroe and Oktibbeha counties, Mississippi, USA, 2018 – 2019. R.A. Nichols et al. Forest Ecology and Management 500 (2021) 119646 3 Fowler et al., 1967). We incorporated plant species in the nutritional constraints model classified as moderate-high deer forage preference based on literature (Warren and Hurst, 1981, Miller and Miller, 1999) along with input from deer biologists with Mississippi Department of Wildlife, Fisheries and Parks (Tables S.4 and S.5). 2.3. Data analysis We used a linear mixed-model analysis of variance using the “ lme4 ” package (Bates et al., 2015) in program R (R core team, 2013) to assess the effects of treatment, year, and treatment × year interactions on deer nutritional quality by forage class, biomass by forage class, and NCC. We classified unburned control and dormant and growing season fire as our treatment fixed effects, species as a random effect for forage quality, and block × year as a random effect for forage biomass and NCC. We clas- sified species as our random effect for forage quality, in order to determine how treatments were affecting quality while taking into ac- count species variation. We did not use block as a random effect because there were no forage quality values for each species at each block. If we found significant treatment main effects, we used the “ lsmeans ” function to compare means within year among treatments and between years within treatments. We used a significance level of α = 0.05 for all contrasts. 3. Results 3.1. Forage quality Crude protein of all forages combined and of forbs, in particular, varied in relation to time since fire (P < 0.05; Fig. 3, Table 1). Crude Fig. 3. Growing-season crude protein by forage class of moderate-high use white-tailed deer forages sampled August-September of 2018 – 2019 in 9 mid-rotation loblolly pine plantations following unburned treatments (Control) and fire treatments during March 2018 (Dormant) and June 2018 (Growing) in Monroe and Oktibbeha counties, Mississippi, USA. Plots are least-square mean and standard error. Table 1 Interactions and main effects of forage quality and biomass by forage class of moderate-high use deer forage species and nutritional carrying capacity (NCC) estimates at a mean dietary level of 14% crude protein (CP) and 0.25% phos- phorus (P). The three treatments included are control (unburned), dormant season fire (March 2018), and growing season fire (June 2018). Treatments were sampled during August-September 2018 and 2019 in 9 mid-rotation loblolly pine plantations in Monroe and Oktibbeha counties, Mississippi, USA. Treatment Yr Treatment × Yr Variable F a P-value F P-value F P-value CP Forb 7.01 0.002 8.99 0.004 6.45 0.003 Vine* 4.09 0.024 5.27 0.027 3.06 0.058 Shrub/Tree 6.25 0.005 11.67 0.002 2.60 0.089 Total 16.99 ≤ 0.001 22.00 ≤ 0.001 10.93 ≤ 0.001 P Forb 2.62 0.081 1.95 0.167 3.71 0.030 Vine* 3.51 0.039 0.48 0.494 0.57 0.572 Shrub/Tree 3.88 0.031 2.06 0.161 0.38 0.686 Total 9.94 ≤ 0.001 1.03 0.311 2.30 0.104 Forage biomass Forb 1.66 0.206 4.75 0.045 8.96 ≤ 0.001 Vine* 18.75 ≤ 0.001 8.99 0.008 0.06 0.941 Shrub/Tree 10.53 ≤ 0.001 2.96 0.105 0.39 0.680 Total 22.33 ≤ 0.001 20.99 ≤ 0.001 3.33 0.048 NCC 14% CP 14.73 ≤ 0.001 6.31 0.023 7.54 0.002 0.25% P 4.73 0.016 1.47 0.243 0.34 0.717 a df determined using Kenward-Roger. * Includes brambles (Rubus spp., Smilax spp.). R.A. Nichols et al. Forest Ecology and Management 500 (2021) 119646 4 protein of all forages averaged 2.8% CP greater in growing season plots as compared to unburned and dormant season plot forages in the first growing season (Fig. 3(d), Table A.1), but the effect was lost during the second growing season. A similar pattern was present in the forb, vine, and woody forage classes. Phosphorus content of all forages, and in particular vines and shrub/ trees, varied by treatment across both years (P < 0.05; Table 1). Phos- phorus of all forages in growing season plots averaged 0.04% P greater than unburned plots and dormant season plots (Fig. 4(d), Table A.1). Vine and shrub/tree P increased in growing season plots by an average of ~0.03 %P compared to unburned and dormant season plots (Fig. 4(b), (c), Table A.1). Forb phosphorus content varied in relation to time since fire (P = 0.030; Table 1). Phosphorus in dormant and growing season plots was 0.07% P greater than in unburned plots in the first growing season and it remained greater in growing season plots during the sec- ond growing season (Fig. 4(a), Table A.1). 3.2. Forage biomass Biomass of all forages combined, and in particular forbs, varied in relation to time since fire (P < 0.05; Table 1). Total biomass of moderate- high use deer forages was greatest in dormant season plots in both growing seasons (Fig. 5(d), Table A.2). Total biomass increased by an average of 1.5 times in unburned and dormant season plots and growing season plots underwent a 3-fold increase in total biomass between growing seasons. Forb biomass in dormant season plots was 7 times greater than growing season plots in the first growing season (Fig. 5(a), Table A.2). Growing season fire forb biomass increased 11-fold between growing seasons and was 2 times greater than unburned and dormant season forb biomass in the second growing season. Vine biomass varied by treatment and year, while shrub/tree biomass varied by treatment across years (P < 0.05; Table 1). Vine biomass in dormant season plots was an average of 1.9 times greater than unburned and growing season plot biomass in both growing sea- sons (Fig. 5(b), Table A.2). Vine biomass in each treatment increased by 1.6 – 2.2 times between growing seasons. Shrub/tree biomass in dormant season plots averaged 3.2 and 8.5 times more biomass than unburned and growing season plots biomass in both growing seasons (Fig. 5(c), Table A.2). 3.3. Nutritional carrying capacity Crude protein-based NCC varied in relation to time since fire (P = 0.002; Table 1). Crude protein NCC in dormant season plots was 2 times greater than unburned plots and 4 times greater than growing season plots in the first growing season (Fig. 6, Table A.3). Crude protein NCC in both fire treatments nearly doubled unburned plots in the second growing season. Growing season plots CP NCC increased between growing seasons by 4-fold. Phosphorus-based NCC varied by treatment across both years (P = 0.016; Table 1). Dormant season P NCC averaged ~49 deer days/ha greater than unburned and growing season P NCC during both growing seasons (Fig. 7, Table A.3). However, growing season P NCC nearly doubled between years and was similar to dormant season one year following fire. 4. Discussion Prescribed fire timing affected nutrient concentrations of moderate- high use deer forages, consistent with our prediction that growing Fig. 4. Growing-season phosphorus by forage class of moderate-high use white-tailed deer forages sampled August-September of 2018 – 2019 in 9 mid-rotation loblolly pine plantations following unburned treatments (Control) and fire treatments during March 2018 (Dormant) and June 2018 (Growing) in Monroe and Oktibbeha counties, Mississippi, USA. Plots are least-square mean and standard error. R.A. Nichols et al. Forest Ecology and Management 500 (2021) 119646 5 Fig. 5. Growing-season biomass (kg/ha) by forage class of moderate-high use white-tailed deer forages sampled August-September of 2018 – 2019 in 9 mid-rotation loblolly pine plantations following unburned treatments (Control) and fire treatments during March 2018 (Dormant) and June 2018 (Growing) in Monroe and Oktibbeha counties, Mississippi, USA. Plots are least-square mean and standard error. Fig. 6. Nutritional carrying capacity (deer days/ha) at a 14% crude protein constraint based on forage species sampled during August-September 2018 and 2019 in 9 mid-rotation loblolly pine plantations following unburned treatments (Control) and fire treatments during March 2018 (Dormant) and June 2018 (Growing) in Monroe and Oktibbeha counties, Mississippi, USA. Plots are least-square mean and standard error. R.A. Nichols et al. Forest Ecology and Management 500 (2021) 119646 6 season fire would increase forage quality during the first growing season (Lewis et al., 1982, Dyke and Darragh, 2007, Eby et al., 2014). Increases in CP and P in growing season plots during the first growing season was most likely due to plants sampled at that time were young. Dormant season fire did not increase forage quality, other than forb P, as compared to unburned controls, which was also consistent with our prediction (Lewis et al., 1982, Dyke and Darragh, 2007, Eby et al., 2014). Dormant season decreased in overall P during the second growing season. However, the greater P of all forages for the growing season fire during the second growing season was contrary to other studies that reported ephemeral gains in nutrition are lost within 3 – 4 months (Lewis et al., 1982). We believe these trends could be contrib- uted to the greater forb biomass produced following growing season fire as opposed to dormant season fire during the second growing season. Forbs are typically higher quality than other forages and may have contributed to the increased overall P levels. This longer lasting increase in P following growing season fire is important in deer management because P positively influences deer plant selection (Lashley et al., 2015c, Dykes et al., 2018) and forages are commonly deficient in P and do not reach minimum P levels required by lactating deer (Blair et al., 1977, Grasman and Hellgren, 1993, Hewitt, 2011). However, the sus- tained increase of 0.07% P following growing season fire increased average P of deer forages to meet minimal lactation requirements. The effects prescribed fire had on forage biomass of moderate-high use deer forages varied based on timing of fire. Greater amounts of biomass during the first growing season for dormant season fire were consistent other studies (Harper, 2007, Lashley et al., 2015a). Growing season fire produced only 74 kg/ha of selected forage biomass during the first growing season. Moreover, biomass of selected forages following dormant season fire increased in both growing seasons following fire over control, and biomass following growing season fires increased but did not equal dormant season fire by the second growing season. The reduction in shrub/tree biomass following growing season fire in this study led us to believe this was due to growing season fire more effectively controlling woody plants. This was consistent with other studies that report fire conducted in the growing season controls woody plant encroachment better than dormant season fire (Drewa et al., 2002, Gruchy et al., 2009). This data suggests that continued growing season fires could cause a decrease in the overall woody component and subsequent increase in forb component, which would ultimately lead to overall increased NCC. The Hobbs and Swift (1985) nutritional constraints model has pre- vented overestimation of NCC because it takes into account minimal dietary requirements and forage availability of deer (Jones et al., 2009). However, calculating NCC with this method should not be considered a true estimate of carrying capacity, but should be considered a relative index reflecting carrying capacity (Edwards et al., 2004, Lashley et al., 2011, Nanney et al., 2018). We selected CP and P as our model con- straints because they are commonly designated as limiting nutrients during summer months for deer (Blair et al., 1977, Grasman and Hellgren, 1993, Hewitt, 2011). We did not include maintenance level estimates in this study because all plants were above the minimum threshold and would have closely resembled forage biomass estimates. Prescribed fire timing had an effect on CP-based NCC, likely due to the effects that prescribed fire timing had on forage quality and biomass (Hobbs and Swift, 1985, Jones et al., 2009). The dormant season fire increase in CP NCC during the first growing season resulted from increased overall forage biomass produced during the growing season. Although growing season fire biomass was lower than dormant season fire biomass in the second growing season, they both improved CP NCC over the unburned control plots during the second growing season. This was similar to previous studies that reported prescribed fire increased CP NCC during the second growing season, or 1-year post fire (Lashley et al., 2011, Nanney et al., 2018). This was caused by the greater amount of forb biomass growing season fire produced. Forbs are a significant portion of a white-tailed deer ’ s summer diet and their likelihood to meet the minimum requirements of CP, especially during harsh environ- mental conditions (Holechek, 1984, Lashley and Harper, 2012, Hewitt, 2011, Nanney et al., 2018), would result in more biomass meeting the mixed diet quality of 14% CP (Lashley et al., 2011). Fig. 7. Nutritional carrying capacity (deer days/ha) at a 0.25% phosphorus constraint based on forage species sampled during August-September 2018 and 2019 in 9 mid-rotation loblolly pine plantations following unburned treatments (Control) and fire treatments during March 2018 (Dormant) and June 2018 (Growing) in Monroe and Oktibbeha counties, Mississippi, USA. Plots are least-square mean and standard error. R.A. Nichols et al. Forest Ecology and Management 500 (2021) 119646 7 Prescribed fire timing also had an effect on P-based NCC. The continued increase in P NCC in dormant season plots was likely due to dormant season fire producing more overall biomass during both growing seasons and few plant species reaching the minimum P threshold for a lactating doe with one fawn. This lead to high variation within NCC estimates among treatments because some sites produced little amounts of biomass and some produced high amounts of biomass that met or exceeded the mean diet quality for P. However, the growing season fire increase in P and biomass during the second growing season caused P NCC to increase. The dormant season fire increase in NCC and growing season fire increase in selected forages CP during the first growing season indicates that timing of fire is an important consideration for land managers. Incorporating multiple fire timings on the landscape will allow deer to utilize high quality forage produced from growing season fire as well as the increased NCC from dormant season fire during lactation. This may be particularly important to provide the opportunity for herbivores to meet their changing nutritional demands through heterogeneity in plant nutritional composition and maturation (Dykes et al., 2020). In our study, it was evident that manipulating fire timing optimizes nutrient availability over time. Therefore, there is a need to incorporate multiple fire timings within fire regimes to adequately optimize forage quality and biomass across years. Incorporating multiple fire timings will make it more logistically feasible for land managers to conduct all planned prescribed burns by extending the burn window into multiple seasons. However, land managers are generally skeptical of burning during the growing season because of smoke management issues and limited burn days (Palik et al., 2002, Ryan et al., 2013). Growing season fire is also associated with direct population-level negative effects on ground nesting birds, but these effects are minimal because fire has the ability to offset direct effects through improved habitat conditions in the future (Cox and Widener, 2008, Kilburg et al., 2014). Variety in timing and spatial scale likely benefit the widest array of species in fire influenced communities (Harper et al., 2016, Mason and Lashley, 2021). Likewise, the impacts of fire timing during sensitive times and over the long-term are likely spatial scale dependent. When forage quality and quantity are at their peaks is dependent on when fire is implemented and pyrodiversity should be an essential consideration when managing for deer habitat and forage quality (Lashley et al., 2015b, Bowman and Legge, 2016). Moreover, deer and other wildlife benefit from the reduction of woody encroachment following growing season fire which can provide more high quality foraging patches during the late summer (Lewis et al., 1982, Gruchy et al., 2009). Integrating more variable fire timing into current regimes will optimize available nutrition and deer NCC on the landscape to help deer better meet their nutritional demands during nutritionally stressed periods and reach their full potential. 5. Conclusions Managing across the landscape using multiple prescribed fire timings can optimize forage quality, forage quantity, and deer NCC during the summer nutritional stress period. Managers can vary fire timing to optimize overall nutrient and forage availability on the landscape. Another important consideration is that the diversity of fire will expand the burn window. Expanding burn windows may be necessary to maintain flammability and arrest transition of forest communities to alternative forest types (Chiodi et al., 2018, Alexander et al., 2021, Kane et al., 2021, McDaniel et al., 2021). We recommend that land managers section properties into smaller burn units and implement prescribed fire during multiple seasons to expand burn opportunities and reduce logistical constraints and promote plant diversity, quality, and quantity to support nutritionally stressed periods for deer. CRediT authorship contribution statement Rainer A. Nichols: Conceptualization, Methodology, Software, Validation, Formal analysis, Investigation, Resources, Data curation, Writing – original draft, Visualization, Project administration. Steve Demarais: Conceptualization, Methodology, Validation, Resources, Writing – review & editing, Visualization, Supervision, Project admin- istration, Funding acquisition. Bronson K. Strickland: Conceptualiza- tion, Methodology, Software, Validation, Writing – review & editing, Visualization, Supervision. Marcus A. Lashley: Conceptualization, Methodology, Validation, Resources, Writing – review & editing, Visu- alization, Supervision, Project administration. Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Acknowledgements We would like to thank the Mississippi Department of Wildlife, Fisheries, and Parks (MDWFP) for financial support through the Federal Aid in Wildlife Restoration Project W-48-50 and the Mississippi State University Forestry and Wildlife Center for funding this research. We would also like to thank the private landowners that allowed us to conduct this research on their properties and technicians/volunteers that assisted in data collection. 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