INFLUENCES OFHUNTING ONMOVEMENTS OF MALEWILD TURKEYSDURING SPRING

John T. GrossWarnell School of Forestry and Natural Resources,

University of Georgia,

Athens, GA 30602, USA

Bradley S. CohenWarnell School of Forestry and Natural Resources,

University of Georgia,

Athens, GA 30602, USA

Bret A. CollierSchool of Renewable Natural Resources, Louisiana State

University Agricultural Center,

Baton Rouge, LA 70803, USA

Michael J. Chamberlain1

Warnell School of Forestry and Natural Resources,

University of Georgia,

Athens, GA 30602, USA

Abstract: Wild turkeys (Meleagris gallopavo; hereafter, turkeys) are an important game species throughout North America,

and interest in turkey hunting continues to increase. Harvest during spring is the primary source of mortality for males, but

little research has addressed how hunting during spring affects movements, space use, and roost site selection of turkeys.

Therefore, we used GPS units attached to male eastern turkeys (M. g. silvestris) and provided to hunters to examine spatial

ecology of turkeys relative to hunting presence in Louisiana. We evaluated effect of hunter presence on daily movement

distances and distances between consecutive roost sites. We also used hunter track logs to create high and low risk gradients

across study landscapes to evaluate how hunter presence impacted weekly core area size and shifts in weekly core area

centers for male turkeys. We monitored 12 male turkeys and found that hunter presence had little effect on male movements;

males increased distances moved by 6% on hunted days (3,976 6 98 m; mean 6 SE; range 1,422–7,751 m, n = 156 days)

compared to non-hunted days (3,741 6 62 m; range 882–7,416, n = 331 days). Distance between consecutive roost sites did

not significantly differ on hunted versus not hunted days. We found that habitation of areas of low risk for hunter presence

tended to result in decreased weekly core area sizes and increased proximity of weekly core area centers to one another.

Habitation of high risk areas for hunter presence increased both core area size and distance between weekly core area centers,

but parameter estimates from these analyses suggested that strength of the observed relationship was weak. We noted that

responses to hunter presence were greatly variable among individuals. Turkeys that survived the hunting season tended to

more often encounter areas of great hunter presence, suggesting a learned antipredator behavior. Managers should consider

altering hunter access across the landscape to create zones of varying risk, thereby affording hunters opportunities to

encounter turkeys with varying levels of experience to hunting.

Proceedings of the National Wild Turkey Symposium 11:259–268

Key words: core area, daily movements, hunter presence, hunting pressure, Louisiana, Meleagris gallopavo, micro-GPS,

roosting, wild turkey.

Wild turkey (Meleagris gallopavo; hereafter, turkey)populations have increased in North America since the1940s, and hunted populations now occur in every stateexcept Alaska (Kennamer 2000, Tapley et al. 2001).Harvest of turkeys during spring is the primary cause ofmortality in adult males (Godwin et al. 1991, Hughes et al.2005, Chamberlain et al. 2012), yet we lack a basicunderstanding of how turkeys respond to hunting activity.Coincident with increasing turkey populations has been an

increase in number of turkey hunters from about 1.69million in 1984–1985 to 2.66 million in 1998–1999, 2.80million in 2009, and 2.49 million in 2014 (Tapley et al.2001, 2011; Eriksen et al. 2015). Nationally, rates of springhunting increased by 22–35% in each 5-year interval during1985 to 2000 (Tapley et al. 2001), increased 7% between

Associate Editor: Healy1 E-mail: mchamberlain@warnell.uga.edu

259

2000 and 2009 (Tapley et al. 2011), but have since declined13% during 2010 to 2014 (Erickson et al. 2015). Althoughhunter attitudes, success rates and demographics have beenstudied (e.g., Palmer et al. 1990, Backs 1995, Thackstonand Holbrook 1995, Little et al. 2001, Casalena et al. 2010),how hunting affects behavior of turkeys is poorlyunderstood.

Everett et al. (1978) used weekly radiotelemetrylocations in Alabama to report that hunting pressure didnot cause shifts in male turkey movement patterns during aspring hunting season. Similarly, in response to a fallhunting season in Florida, Williams et al. (1978) reportedthat most Osceola turkeys (M. g. osceola) maintained theirpre-season ranges and roosting sites, with some turkeysactually decreasing movements. Research on response tohunting for other game species has demonstrated that somespecies change or increase movements both spatially andtemporally (Root et al. 1988, Kilgo et al. 1998), whereasother species do not typically alter patterns of space use(Hodges et al. 2000, Karns et al. 2012).

Daily movements of turkeys have not been wellstudied, as most research has focused on seasonalmovements, dispersal, and space use (Kelley et al. 1988,Holdstock et al. 2006). Daily distances moved by maleshave been found to be greater in spring than other seasons(Godwin et al. 1994) and long distance daily movementscould influence individual survival and management goals.Such daily movements are often centered on core areas ofuse presumably containing critical resources important forreproduction and survival (Asensio et al. 2012). It isplausible to assume that turkeys could alter size of theircore area, their daily movement within these core areas, orpotentially shift their core areas in response to hunting anddisturbance. Beyond core areas and daily movementswithin, selection of roost sites also could influence survivalas roosts provide protection from inclement weather andpredation (Porter 1978, Kilpatrick et al. 1988). Huntingturkeys often puts hunters close to roost sites well beforeturkeys leave a roost, and this disturbance could influenceroost site selection. Roosting behavior in turkeys is poorlyunderstood relative to other facets of their ecology, and alsocould be affected by hunter disturbance.

Previous studies assessing space use and movements ofmale turkeys have used locations collected via VHFradiotransmitters and often contained great locational error(White and Garrott 1986, Thogmartin 2001, Montgomeryet al. 2011). However, advent of GPS units designedspecifically for turkeys has increased our capability togather data that are more spatially and temporally accurate,and free of researcher biases (Guthrie et al. 2011). Forexample, Collier and Chamberlain (2010) reported that amale turkey fitted with a GPS unit moved approximately1,000 m and shifted its roost location on the opening day ofthe youth hunting season in south Louisiana. This malethen exhibited short-distance linear movements for severaldays, until moving approximately 2,000 m on opening dayof a 50-hunter lottery hunt. Given our lack of understandingof how ephemeral presence of hunters on the landscapeaffects spatial ecology of turkeys, our objectives were touse GPS technology to describe male turkey responses tospring hunting. Specifically, by monitoring hunter andturkey behavior simultaneously with GPS technology, we

evaluated how hunting affected daily movements, place-ment and sizes of core use areas, and roost site selection ofmale turkeys during spring hunting season.

STUDY AREA

We conducted our research on 3 study sites during2012–2013. Our primary study area was the 2,390-haTunica Hills Wildlife Management Area (WMA) located inWest Feliciana Parish, Louisiana. Tunica Hills was ownedand operated by the Louisiana Department of Wildlife andFisheries, and was located at the southernmost edge of theloess blufflands. Tunica Hills and surrounding private landswere composed of dissected uplands characterized by steepbluffs, ravines and rugged hills. Forest types of TunicaHills were upland hardwoods with the primary soil typebeing wind-deposited silt loams from the Mississippi River.Common overstory species included American beech(Fagus grandifolia), oaks (Quercus spp.), hickories (Caryaspp.), eastern hophornbeam (Ostrya virginiana), yellow-poplar (Liriodendron tulipifera), red maple (Acer rubrum),loblolly pine (Pinus taeda), and eastern red cedar(Juniperus virginiana). Understory shrubs and plantsincluded oak leaf hydrangea (Hydrangea quercifolia),two-wing silverbell (Halesia diptera), pawpaw (Asiminatriloba), muscadine grape (Vitis rotundifolia), floweringdogwood (Cornus florida), sweetleaf (Symplocos tinctoria),blackberry (Rubus spp.), and switchcane (Arundinariagigantea).

Turkey hunting was regulated with a season structurethat allowed for a lottery system and a one-week hunt opento the public. Turkey season began with a one-day youthhunt on the third Saturday in March of every year, followedby 3 weekends (Saturday and Sunday only) of lotteryhunting, in which only 15 participants were allowed tohunt. After the last Sunday of lottery hunting, the WMAwas open to the public for 7 days.

Mosher Hill Hunting Club (hereafter Mosher) was a1,700-ha property in Washington Parish owned andmanaged by Weyerhaeuser Company and leased to aprivate hunting club. Mosher had 100 club members, ofwhich 10 were spring turkey hunters. Turkey season beganwith a 2-day youth hunt opening on 16 March 2013,followed by a 4-week open season that began the followingweekend and concluded on 14 April. Most hunting activitywas concentrated on weekends, although hunting during theweek did occur. Mosher was comprised mostly of loblollypine (Pinus taeda) and was intensively managed for pinesawtimber production. The area also contained small riverdrainages and low areas that consisted mostly of hardwoodspecies including water oak (Q. nigra), green ash (Fraxinuspennsylvanica), sweet bay (Magnolia virginiana), southernmagnolia (Magnolia grandiflora), wild azalea (Rhododen-dron canescens), baldcypress (Taxodium distichum), watertupelo (Nyssa aquatica), and red bay (Persea borbonia).

Located in West Baton Rouge Parish, Louisiana,Double D hunting club (hereafter Double D), was a 688-ha privately owned hunting club. Double D was approx-imately 5 km north of Interstate 10 in the floodplainsbetween the Mississippi and Atchafalaya rivers. The areahad 13 club members, 4 of whom were turkey hunters.Turkey season began with a 2-day youth hunt opening on

260 Harvest Management and Hunting

16 March 2013, followed by a 5-week open season thatbegan the following weekend and concluded on 21 April.Most hunting activity was concentrated on weekends,although hunting during the week did occur. The propertycontained bottomland hardwoods and a mix of roadways,gas pipelines and food plots. Due to a closed canopy andprolonged seasonal flooding, much of the mid andunderstory was sparse. Overstory species included wateroak, nuttall oak (Q. nuttalii), overcup oak (Q. lyrata),eastern cottonwood (Populus deltoides), American syca-more (Platanus occidentalis), bitter pecan (Carya xlecontei), water hickory (C. aquatica), sweetgum (Liquid-ambar styraciflua), sugarberry (Celtis laevigata), and ash(Fraxinus spp.). Midstory and understory species includedred mulberry (Morus rubra), boxelder (Acer negundo), redmaple, roughleaf dogwood (C. drumondii), trumpet creeper(Bignonia capreolata), dewberry (Rubus spp.), and poisonivy (Toxicodendron radicans).

METHODS

We captured male turkeys eastern turkeys (M.gallopavo silvestris; hereafter, turkeys) using cannon netsduring January–March 2012 on Tunica Hills and adjacentprivate lands. Due to a lack of trapping success on TunicaHills in 2013, we expanded trapping efforts to Mosher andDouble D hunting clubs that year. Study sites were over100 km from each other; we considered each as separatelocations, but each received low hunting pressure (150þ haper hunter). Once captured, we attached GPS units to maleturkeys using 3-mm shock cord backpack style (Wilson andNorman 1995). We also fitted turkeys with aluminum rivetbands on their right tarsus. We estimated age (adult orjuvenile) based on development and barring of 9th and 10th

primary feathers (Pelham and Dickson 1992). The Univer-sity of Georgia Institutional Animal Care and UseCommittee approved capture and handling protocols(Permit A2011 07-003-R1).

We fitted all turkeys with GPS units designed andproduced by either Sirtrack Wildlife Tracking Solutions(Sirtrack Wildlife Tracking Solutions, Havelock North,New Zealand) or MiniTrack Backpack GPS units (BiotrackLimited, Wareham, Dorset, United Kingdom). Sirtrackunits were approximately 10 cm 3 4 cm, were programmedbefore deployment, and had to be recovered to retrievedata. Units produced by Biotrack measured approximately7.5 cm 3 2.5 cm and were capable of having data remotelydownloaded. All units housed both GPS and VHFcomponents, allowing us to triangulate turkeys once perweek to monitor survival. We programmed GPS units torecord one location at noon and one location at midnighteach day from time of capture through March 10 of thesame year. On March 10, we programmed GPS units toswitch to a greater intensity schedule, collecting onelocation every 15 minutes during daylight hours and onelocation at midnight each day.

We continued this schedule for 50 days, whichcoincided with all hunting activity. After the 50 days wascomplete, units began recording one location at noon onceper week to provide coarse location data and to maintainbattery life so that units could collect data in 2 consecutivesprings. We programmed units to repeat this schedule

beginning on March 10 of the following year if the turkeywas still being monitored.

The Louisiana Department of Wildlife and Fisheriesrequired all hunters on Tunica Hills to stop at designatedcheck-in stations upon entry. We briefly discussed ourstudy objectives with each hunter, before providing themwith a handheld Garmin eTrex GPS unit. We pre-programmed each unit to collect one location every 30sec for the duration of the hunt. We turned each unit onbefore giving it to the hunter and asked them to carry itthroughout duration of his or her hunt. Each GPS unit wasequipped with a belt clip, but most hunters chose to placeunits in an extra pocket or a backpack. We collected unitsfrom hunters upon exiting the WMA, at which time wedownloaded data and cleared units for re-deployment.During the 2012 hunting season, it was not mandatory forhunters to carry GPS units, but only 3 hunters refused toparticipate in our study. During the 2013 hunting season,the Louisiana Department of Wildlife and Fisheries madeparticipation mandatory.

On Mosher and Double D, we asked hunters tovoluntarily pick up a GPS unit from designated check-instations before every hunt. We programmed these GPSunits similarly to those deployed on Tunica Hills. Webelieve that we captured most (.90%) hunting events onboth private lands. We then retrieved units twice weekly,downloaded data from them, and cleared them for re-deployment.

We evaluated how hunting influenced linear distance(m) traveled per day, distance (m) between consecutiveroost sites, shifts in weekly core area centers (m), andweekly core area size (ha). We limited our analysis periodfrom 1 March to 30 April during 2012 and 2013 to coincidewith peak hunting and breeding activity on our sites.

To evaluate how daily linear distances moved wereaffected by hunter presence, we first calculated lineardistance (m) traveled per day using the XY to Line tool inArcGIS 10.0 (ESRI 2011). We then used each recordedlocation from that day to create a total daily movement pathand measured total linear distance. To evaluate if huntinginfluenced roost site locations, we calculated distances (m)between nightly roost locations during spring huntingseason. Using GPS locations collected every night atmidnight, we measured distance between consecutive roostlocations in ArcGIS 10.0 (ESRI 2011). We then applied 2-sample t-tests to compare differences in daily movementsand distances between consecutive roost sites betweenhunted and non-hunted days.

To determine if hunter presence affected weekly coreuse areas, we first grouped turkey locations into weekscorresponding to the Tunica Hills hunting season. We usedprogram R version 3.1.0 (R Development Core Team2013), and package adehabitatHR (Calenge 2006) tocalculate kernel density core-use areas (50%) for eachturkey for each week. Once we calculated 50% core areas,we calculated centroids for each week and measured shiftin location of each centroid over consecutive weeks.

We used hunter track logs to create daily risk gradientsof hunter presence. We used these ‘‘risk zones’’ todetermine likelihood of each turkey encountering a hunterduring each day of hunting season. Our 2 risk gradientswere based on visual and auditory detection; visual

Hunting and Male Turkey Movements � Gross et al. 261

detection of a predator was a high risk situation, andauditory detection of a predator was a low risk situation.Although these landscapes varied both in midstory densityand topography, we believed that 100 m was a reasonabledistance a turkey may have been able to detect a hunterwith visual perception in these hardwood and pine-dominated landscapes. Heffner and Heffner (1992) notedthat the turkey’s auditory sensitivity is less than that of ahuman. Likewise, the inverse square law states that, in anenvironment with no obstructions, sound intensity isinversely proportional to distance squared from the soundsource. Thus, we estimated that a sound at an intensity of100 dB (considered a loud sound) could be detected by aturkey as far as 300 m in typical conditions across these 3landscapes. Therefore, we designated high risk hunterpresence zones as any area within 100 m of a recordedhunter location for that day and low risk hunter presencezones as anything between 100 m and 300 m. To calculatethese risk zones, we combined all hunter data from one dayinto a single GIS file and then created a raster layer usingthe Euclidean Distance tool in ArcMap (ESRI 2011). Wereclassified this raster layer into high and low risk zonesbased on the 100 m and 300 m definitions. To calculatehunter presence, we separated all recorded turkey locationsby day and projected this onto calculated hunter presencemaps. Once we displayed daily risk zones and associatedturkey locations, we used the Extract Values to Points toolin ArcMap to sum number of points within high and lowhunter presence zones. We then summed these values foreach week.

We conducted generalized linear mixed modeling(GLMM) analyses using the lme4 (Bates et al. 2011) andLMER Convenience Functions (Tremblay 2011) to inves-tigate if hunter presence affected core area size and shifts.The GLMMs allowed us to incorporate a flexiblecovariance structure into the modeling framework, result-ing in better estimates of variability than standardgeneralized linear models (Clayton and Kaldor 1987,Breslow and Clayton 1993). We used a log link withPoisson error term in these analyses, and fitted modelsusing Laplace approximation. The dependent variableswere the aforementioned weekly core area shifts and corearea size. Fixed response variables were weekly counts ofhigh and low risk events. To account for assumedindividual behavioral characteristics and repeated measuresof each turkey, we treated each turkey as a random effect.Although our samples were from 3 study sites, we ignoredthis factor as a random effect because of insufficientreplication. To examine if increasing risk events wereassociated with increased hunter-induced mortality, wedetermined mean and standard error of number of weeklyhigh and low risk encounters for turkeys that survivedhunting season and turkeys that were harvested by hunters.We conducted all analyses using program R version 3.1.0(R Development Core Team 2013) using an a priori a =0.05.

RESULTS

We captured 17 adult and 2 juvenile males during2012–2013 and examined influences of hunting onmovements of 11 adult and 1 juvenile turkey. Of these 12

turkeys, we captured 7 adults from Tunica Hills in 2012, 1juvenile from Double D in 2013, and 4 adults from Mosherin 2013. Due to small sample sizes and having only 1juvenile turkey in our data set, we pooled all turkeys for ouranalyses. We obtained an average of 1,964 locations perturkey during our study period. We observed an 8%increase (t11 =�2.16, P = 0.05) in daily distances traveledfor hunted days (4,022 6 98 m; mean 6 SE; range 1,422–7,751 m, n = 156 days) compared to non-hunted days(3,740 6 62 m; range 882–7,416, n = 331 days). Distancebetween consecutive roost sites did not differ (18%, t11 =�1.488, P = 0.165) on hunted days (947 6 69 m; range 7–4,350; n = 151 roost events) versus non-hunted days (7986 36 m; range 3–3,548; n = 363 roost events).

We found that low risk hunter presence tended to resultin decreased weekly core area sizes (b =�0.010; Table 1)and increased proximity of weekly core area centers to oneanother (b = �0.010; Table 2). High risk hunter presenceincreased both core area size (b = 0.008; Table 1) anddistance between weekly core area centers (b , 0.001;Table 2). Average weekly core area sizes ranged from 454ha to 120 ha and average shift in weekly core area rangedfrom 437 m to 1,234 m. Individual turkey behavior wasvariable with regards to core area size (parameter estimatesof �1.92 to 0.89) and shift in weekly core area centers(parameter estimates of �1.19 to 1.4; Fig, 2). To illustratedegree of variability, one male moved beyond the TunicaHills WMA boundary on opening day of hunting seasonand never returned (Fig. 3), whereas other males main-tained core areas comprised of .50% of total area in whatwe delineated as high risk hunter presence areas. Hunterskilled 7 of 12 males. Turkeys harvested by hunters tendedto have fewer high risk encounters each week (10.82 6

3.16) than those that survived hunting season (31.00 6

10.54). Contrastingly, low risk encounters tended to occurmore for harvested males (24.39 6 13.00) than forsurviving males (15.37 6 7.05). This finding suggests thatperhaps turkeys experiencing less pressure are easier toharvest.

Table 1. Parameter estimates (log scale) quantifying effect ofnumber of different levels of hunter risk zones on weekly core

area size (ha) of 12 male eastern wild turkeys in Louisiana duringspring turkey hunting seasons of 2012 and 2013. High risk

hunter presence zones were areas within 100 m of a recorded

hunter location for that day and low risk hunter presence wasanything between 100 m and 300 m. Standard errors (SE), z

values, and probabilities that a coefficient differs from 0 are alsopresented.

CovariateEstimate

(b)Coefficient

(SE) z value Pr(.jzj)

Intercept 4.18 0.22 19.34 ,0.01

High Risk Hunter

Presence 0.0087 7.01e-4 12.36 ,0.01

Low Risk Hunter

Presence –0.0100 9.17e-4 –10.91 ,0.01

Turkey 0.55a N/A N/A N/A

a Turkey was considered a random effect in the model. Thus, it is a

variance estimate.

262 Harvest Management and Hunting

DISCUSSION

We found that daily distances moved by turkeys inLouisiana were greater on hunted days than on non-hunteddays. Although daily distances moved were statisticallygreater on hunted versus non-hunted days, turkeys onlymoved about 8% more on hunted days and thesedifferences may not reflect biological relevance. Wesuspect that this finding is simply a consequence of theresolution and volume of data we collected. As such, ourresults seem to support findings of Everett et al. (1978) andWilliams et al. (1978), who reported no changes in malemovements due to hunting pressure. Karns et al. (2012)found that white-tailed deer (Odocoileus virginianus)significantly reduced their daily movements during ahunting season when compared to other seasons. Ourapproach provided finer resolution data relative to howhunter presence affected animal movements because weevaluated differences in movements when hunters werepresent or recently present, not just during an entire huntingperiod. It is important to note that our methodology inassigning risk zones means these counts of hunter presencedo not necessarily mean that a turkey and a hunter werewithin these zones at the same time. Thus, our results maynot necessarily equate to direct proximity of turkeys andhunters to each other at a single point in time, but ratherrepresent how often turkeys spent time in areas with recent

Table 2. Parameter estimates (log scale) quantifying effect of

number of locations within different levels of hunter risk zones onweekly core area shifts (m) of 12 male eastern wild turkeys in

Louisiana during the spring turkey hunting seasons of 2012 and2013. High risk hunter presence zones were areas within 100 m

of a recorded hunter location for that day and low risk hunterpresence was anything between 100 m and 300 m. Standard

errors (SE), z values, and probabilities that a coefficient differsfrom 0 are also presented.

CovariateEstimate

(b)Coefficient

(SE) z value Pr(.jzj)

Intercept 6.76 0.19 35.11 ,0.01

High Risk Hunter

Presence 0.0004 2.14e-4 1.77 0.08

Low Risk Hunter

Presence –0.0104 3.00e-4 –34.86 ,0.01

Turkey 0.44a N/A N/A N/A

a Turkey was considered a random effect in the model. Thus, it is a

variance estimate.

Figure 1. Euclidean distance calculated from hunter track logs

and associated male eastern wild turkey locations at Tunica HillsWMA, Louisiana, from 2013. Hunter track logs (red dots) were

used to create Euclidean distance calculations of high risk zones(100 m, orange buffer) and low risk zones (300 m, yellow buffer)

to calculate high and low risk hunter presence. Blue dotsrepresent turkey locations for the same day as hunter track logs.

Turkey locations within the orange buffer were classified as highrisk hunter presence and locations in the yellow buffers were

classified as low risk hunter presence.

Figure 2. Estimated random effect values of each individual maleeastern wild turkey for weekly core area size (a) and shift in

weekly core area (50%) centers (b) in response to hunterpresence. Random effects were assumed to be normally

distributed with a mean of zero.

Hunting and Male Turkey Movements � Gross et al. 263

hunter use. We offer that these distinctions are importantbecause many variables (and the way they are measured)can result in researchers finding movement patterns to shifttemporally. Karns et al. (2012) noted that although theysaw a decrease in movements during periods when white-tailed deer were hunted, this period also coincided with thepost-breeding period, which may have caused decreasedmovement.

Consecutive roost sites were 18% farther away fromone another on hunted days when compared to non-hunteddays. In Colorado, Hoffman (1991) found male Merriam’s(M. g. merriami) turkeys used consecutive roosts thataveraged over 1,000 m apart and only used a previous roostsite 19% of the time. Our reported estimates of 909 m and766 m between consecutive roosts for hunted and non-hunted days are similar to the findings of Hoffman (1991).Although not statistically significant, we speculate that thisdifference may be biologically relevant as males mayincrease their distance between consecutive roosts ifhunters were present when they left their morning roostsites, or if they were disturbed in areas surrounding a roost.

We observed that low risk hunter presence decreased

core area size and decreased proximity of weekly core areasto one another, whereas high risk hunter presence increasedcore area size and increased distances between weekly coreareas. While observed effect sizes were relatively small, wespeculate that perhaps low risk or indirect hunter presencemay cause turkeys to reduce their core areas, but not shiftthese areas as perceived risk from hunting does not warrantsuch a response. Conversely, when hunter presenceincreases (high risk), turkeys may be more likely toincrease their core area size and shift these areas.Regardless, it is important to note that responses to hunterpresence varied widely among individual turkeys, suggest-ing that responses may have been based on previousexperiences with predation risk or influenced by variableswe did not measure.

Our findings generally paralleled those published fromstudies designed to assess interactions between predatorsand their prey. Ydenberg and Dill (1986) noted that preymade optimal fleeing decisions that balanced chance ofcapture with the trade-offs of increased energy demand.Likewise, probability of fleeing or shifting movementsbeing related to severity of the approach by a predator isplausible, in that an approach that is more direct and incloser proximity to an animal may be more likely to causeit to flee (Burger and Gochfeld 1998, Cooper 1998, Fridand Dill 2002). Dill (1974) hypothesized that flightinitiation distances, or fleeing probability, would increasewhen predators were larger or increased speed in whichthey approached their prey. Similarly, flight initiationdistances should increase when distance to the nearestrefuge increases because risk of capture would alsoincrease, and flight initiation distances would increasewhen resources are low (Frid and Dill 2002). Our resultsconcur with these predictions, and male turkeys appearedmore likely to change their movement patterns when facedwith high risk hunter presence.

Male turkeys move more in spring than in otherseasons (Kelley et al. 1988, Godwin et al. 1994), primarilya result of breeding behaviors. Female movements,courtship and mating behaviors, and social pressurebetween dominant and subordinate males can influencemovements of males in spring (Shields 1987, Hurst et al.1991, Holdstock et al. 2006). Our results show that huntinghad minor effects on space use of males during spring.Further, timing of hunting season, coinciding with peak inbreeding behavior, may have mitigated effects of hunterpresence because males were more concerned withpursuing breeding opportunities than avoiding predators.

Turkeys in our study that spent a smaller proportion oftheir time in high risk zones tended to be more frequentlyharvested. This suggests that male turkeys that frequentlyencounter hunters may learn behaviors that mitigate theirrisk to harvest. Although speculative due to small samplesize, turkeys in greatly pressured areas may not be moresusceptible to harvest. Hunters in one of our study sites(Tunica Hills) relied primarily on roads, with 50% oflocations occurring within 18 m of a trail or road and ononly 2.9% of the WMA (Gross et al. 2014), yet none of ourturkeys whose home range centered in this area wereharvested by hunters. It is possible that male turkeys in highrisk areas learn to associate cues such as hunter-made callswith danger, and may make subtle changes in their

Figure 3. A depiction of what we speculate to be a male eastern

wild turkey avoiding hunter activity by leaving the managementarea on opening day of hunting season at Tunica Hills WMA,

Louisiana in 2012. Yellow points indicate the turkey’s locationsfrom 15 February to 16 March 2012, the red line with arrows

shows the turkey’s movement path on 17 March 2012 (openingday of hunting season), and blue points indicate the turkey’s

locations from 18 March to 7 April when the turkey was killed.Black lines are the hunter track logs recorded with handheld

GPS units on opening day.

264 Harvest Management and Hunting

movements to avoid harvest. Our techniques would nothave been able to identify these subtle avoidancemovements and examining these behaviors may bewarranted. Conversely, our findings suggest that turkeysexperiencing less pressure may be easier to harvest.

We found a wide variation in response to huntingamong individual turkeys. While some males were killedby hunters (Fig. 4), some males seemed to avoid hunterinteraction by fleeing (Fig, 5). Miller et al. (2001) andGrisham et al. (2008) reported a great degree of variabilityin spatial fidelity by males during spring. Grisham et al.(2008) reported that half of the males in their study shiftedtheir space use, but they were unable to detect a pattern atthe population level. We suggest that lack of a consistentpattern among individuals in our study signals that otherfactors, such has breeding behavior, female movements,dominance status, and habitat characteristics, were moreinfluential on male movements than was hunter presence.

MANAGEMENT IMPLICATIONS

Although likely variable across landscapes andsubspecies of turkey, effects of hunting on spatial ecologyand movement of turkeys appear minor. Our findingssuggest that it may be easier to harvest turkeys that

experience less hunting pressure, whereas turkeys experi-encing greater pressure may learn to avoid hunters.Clearly, managers are tasked with optimizing the balancebetween hunter access and hunt quality. We suggest thathunter access and its subsequent effects on individualturkey behavior across managed lands may exert stronginfluences on susceptibility of turkeys to harvest. Superfi-cially, our findings suggest that encouraging hunters toavoid high pressure zones should translate into thosehunters encountering less experienced turkeys, therebyrealizing increased hunting quality and success. However,there is potential at some point that heavily hunted areasmay hold a population of male turkeys that are allrelatively experienced and difficult to harvest. Therefore,managers should consider altering hunter access across thelandscape to create zones of varying risk, thereby affordinghunters opportunities to encounter turkeys with varyinglevels of experience to hunting. For example, managerscould partition land into different management zones opento hunting at different times during the spring season. In ascenario of unacceptably great initial turkey harvest earlyin the season, managers could keep the same managementzone open and leave others closed, which would placehunters in areas where greatly pressured turkeys wereexperienced and less susceptible to harvest. Alternatively,

Figure 4. Recorded movement paths of 1 hunter and 1 maleeastern wild turkey in West Feliciana Parish, Louisiana in 2013.

The yellow line depicts the movement path of a male wild turkey

leaving his roost (first red arrow) and moving approximately1,200 m in 1 hour where he was killed by a stationary turkey

hunter (red star). The hunter path is depicted by red locations.

Figure 5. Movement paths recorded via GPS for 1 hunter and 1male eastern wild turkey in West Feliciana Parish, Louisiana in

2013. The yellow line depicts movement path of a male wildturkey; the blue points are a hunter track log. The red star

indicates where the hunter and turkey met. After the turkeyinteracts with the hunter, he moves .3,000m before roosting

that night.

Hunting and Male Turkey Movements � Gross et al. 265

in areas where male harvest is below target levels or wherehunter success in early portions of the season was less,managers could open new zones to hunting and allowhunters to interact with less experienced turkeys. We offerthat studies examining learned antipredator responses ofturkeys to hunters, and whether these behaviors affecthunter success, are warranted.

ACKNOWLEDGMENTS

We appreciate funding provided by the LouisianaDepartment of Wildlife and Fisheries, the National WildTurkey Federation (NWTF), the Louisiana Chapter ofNWTF, and the Warnell School of Forestry and NaturalResources at the University of Georgia. We thank J.Stafford and J. Wood for field assistance. We thankWeyerhaeuser Company and Double D Hunting Club foraccess to their property during this study.

LITERATURE CITED

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John T. Gross is an environmental consultant for the oil and gas

industry in Houston, Texas. He received his B.S. in Natural

Resource Management and Ecology from Louisiana State Univer-

sity and his M.S. in Forest Resources from the University of

Georgia, where he studied the movement ecology of male wild

turkeys in the southeastern United States.

Bret A. Collier is an Assistant Professor in the School of Renewable

Natural Resources at Louisiana State University. Bret’s research

focus is wildlife population dynamics and development of statistical

methods for wildlife biologists, although he has been known to

delve into a variety of wildlife-related topics. He has been actively

conducting research on wild turkey demography and spatial ecology

for the past 12 years. Bret and his wife, Reagan, have a daughter,

Kennedy, and he is both a hunter and landowner.

Hunting and Male Turkey Movements � Gross et al. 267

Bradley S. Cohen is a post-doctoral researcher at the University of

Georgia. He received his bachelor’s degrees in Biology and

Psychology from the State University of New York at Geneseo,

and his M.S. and Ph.D. degrees in Wildlife Ecology and

Management from the University of Georgia. His research focuses

on the perceptual and behavioral ecology of game animals.

Michael J. Chamberlain is a Professor of Wildlife at the Warnell

School of Forestry and Natural Resources at the University of

Georgia. Mike received his B.S. degree from Virginia Tech, and his

M.S. and Ph.D. degrees from Mississippi State University. Mike’s

research interests are broad, but he focuses much effort into

evaluating relationships between wildlife and their habitats. He has

conducted research on wild turkeys for the past 20 years. Mike is a

dedicated hunter and dad, and enjoys spending time outdoors

regardless of the pursuit.

268 Harvest Management and Hunting