Research Article Survival and Productivity of Eastern Wild Turkey Females in Contrasting Landscapes in Wisconsin CHRISTOPHER D. POLLENTIER, 1,2 Department of Forest and Wildlife Ecology, University of Wisconsin-Madison, 226 Russell Labs, 1630 Linden Drive, Madison, WI 53706, USA R. SCOTT LUTZ, Department of Forest and Wildlife Ecology, University of Wisconsin-Madison, 226 Russell Labs, 1630 Linden Drive, Madison, WI 53706, USA SCOTT D. HULL, Bureau of Science Services, Wisconsin Department of Natural Resources, 2801 Progress Road, Madison, WI 53716, USA ABSTRACT Eastern wild turkeys ( Meleagris gallopavo silvestris ) have colonized the entire state of Wisconsin, USA since being successfully reintroduced in the 1970s. Recently, conservation groups have expressed concerns regarding habitat quantity and quality in association with population plateaus and declines in local abundance in some portions of the state. Estimates of survival and productivity are needed to identify the current population status and for drafting effective management plans. We evaluated female eastern wild turkey demographic parameters and the relationship of these parameters to landscape composition between townships that were forest-dominated and those that were primarily open (e.g., cropland, pasture) in southwest and west-central Wisconsin. We radiomarked 129 female eastern wild turkeys during a 2-year field study (2010–2012). Seasonal variation and landscape effects on survival were important components in our top models, which accounted for > 98% of the relative support. Seasonal survival ( SE) was lowest during spring (15 Mar–18 Jul) across both landscapes (0.672 0.039). The model- averaged estimate of annual (15 Mar–14 Mar) female survival across all study townships was 0.515 0.035 and survival estimates were lower in forested landscapes (2010 ¼ 0.450 0.075, 2011 ¼ 0.279 0.094) than in open landscapes (2010 ¼ 0.707 0.053, 2011 ¼ 0.651 0.045). Predation accounted for 77.8% of all mortalities and was highest during spring; predator-related mortalities were more frequent in forested townships ( P ¼ 0.027). Nest survival was similar in forested landscapes ( ^ u ¼ 0 : 273 0 : 08) and open landscapes ( ^ u ¼ 0 : 201 0 : 07). Additionally, we estimated apparent nesting success to be 0.256 0.04 and it was not different between landscapes ( P ¼ 0.573). Poult survival was lower in forested landscapes compared to open landscapes at 4 weeks post-hatch ( P ¼ 0.008). Differences in eastern wild turkey population demographics may exist in separate local populations that are in close albeit different types of landscapes. Management of eastern wild turkeys that enhances female and poult survival during the nesting and brood-rearing periods can greatly influence population growth, particularly in forest-dominant landscapes. Ó 2014 The Wildlife Society. KEY WORDS eastern wild turkey, forest cover, land cover composition, Meleagris gallopavo silvestris , open cover, productivity, survival, township, Wisconsin. Eastern wild turkeys ( Meleagris gallopavo silvestris ; hereafter, turkeys) were historically common and occupied much of southern Wisconsin, USA, but populations were extirpated by the late 1800s through unregulated market hunting and widespread clearing of forest land (Kubisiak et al. 2001). Repeated stocking attempts with game farm turkeys through the mid-1960s proved unsuccessful; consequently, limited spring male-only hunts were discontinued in the early 1970s (Kubisiak et al. 2001). The Wisconsin Department of Natural Resources (WDNR) renewed reintroductions of turkeys throughout southern Wisconsin from 1976 to 1985 with wild-trapped turkeys from Missouri that proved successful. Additional intrastate translocations were com- pleted by the mid-1990s and current population estimates range between 360,000 and 450,000 birds (Tapley et al. 2011). With restoration efforts in Wisconsin complete, focus has shifted to effective management of local populations and the development of sound harvest strategies. Even though turkeys are established throughout much of the state, questions remain about possible long-term changes in the quantity and quality of habitat that could limit local population growth of turkeys. Given the dynamic nature of agricultural and forest-land management practices, we addressed how and if turkey vital rates varied across landscapes. We took the approach of selecting landscapes Received: 24 July 2013; Accepted: 13 May 2014 Published: 12 July 2014 1 E-mail: christopher.pollentier@wisconsin.gov 2 Present address: Bureau of Science Services, Wisconsin Department of Natural Resources, 2801 Progress Road, Madison, WI 53716, USA The Journal of Wildlife Management 78(6):985–996; 2014; DOI: 10.1002/jwmg.749 Pollentier et al. Wild Turkey Ecology in Contrasting Landscapes 985 with different characteristics, and evaluated survival and nesting success of turkeys in these landscapes to direct management and ensure limited conservation funds are used most efficiently. Land cover characteristics often influence turkey abundance and distribution, and studies have suggested that homogeneous landscapes dominated by 1 land cover type (i.e., agriculture or forest) may be less suitable for turkeys (Gustafson and Parker 1994, Pollentier 2012), whereas landscapes with high heterogeneity and intersper- sion of forests and agriculture can be beneficial (Glennon and Porter 1999). Based on previous work on differences in the abundance of male turkeys between forested and open landscapes in west-central Wisconsin (Lechmaier 2008), we were interested in evaluating female survival and productivity in these landscapes. Extensive information exists regarding eastern wild turkey population dynamics across their range (Vangilder 1992, Palmer et al. 1993, Roberts et al. 1995, Vangilder et al. 2001, Moore et al. 2010) and in Wisconsin (Wright et al. 1996, Paisley et al. 1998). Such past work has identified differences in demographic parameters across a spectrum of landscapes and ecoregions. Although demographic comparisons most often are made across studies, factors such as weather variables (Porter et al. 1983, Roberts et al. 1995), harvest strategy (Healy and Powell 1999, Tapley et al. 2011), or spatial scale of observation (Wiens 1989) may complicate comparisons and obscure differences. Our objective was to compare female turkey survival and productivity concurrently among study sites that were within close proximity (i.e., 100 km) but differed in cover type. Our goal was to identify potential differences in vial rates of female turkeys between predominately forested landscapes and open landscapes with moderate forest cover to guide management strategies used to conserve wild turkey populations. STUDY AREA We investigated female turkey survival and productivity in contrasting pairs of landscapes with different proportions of forest cover and open land in rural southwest and west- central Wisconsin (Fig. 1) within the Central Sand Plains and Western Coulees and Ridges ecoregions (Kreitinger et al. 2012). Paired study sites were located in 4 Public Land Survey System townships (9,300 ha each; hereafter, town- ship): Fairchild (Eau Claire County; 44 8 38 0 N, 90 8 59 0 W), Hale (Trempealeau County; 44 8 28 0 N, 91 8 18 0 W), Stark (Vernon County; 43 8 35 0 N, 90 8 36 0 W), and Westford (Rich- land County; 43 8 30 0 N, 90 8 14 0 W). Topography was primari- ly rugged with steep slopes and deep valleys, and elevation ranged from 270 to 379 m. Dairy farming and row crop agriculture (e.g., corn, soybeans, alfalfa) were principal land uses. Forested areas primarily consisted of oaks ( Quercus spp.), hickories ( Carya spp.), maple ( Acer spp.), aspen ( Populus spp.), and pine ( Pinus spp.). The region encompass- ing all 4 study townships had a mean annual rainfall of 85.3 cm and an average annual snowfall of 109.4 cm with sporadic periods of deep accumulating snow ( > 25 cm) during winter. The coldest month, January, averaged temperatures of 11.1 8 C, and the warmest month, July, averaged 21.2 8 C (Wisconsin State Climatology Office 2012). The majority of land in all 4 study townships was privately held (combined approx. 32,044 ha, 86%). State- and county-managed lands, county forests, and easements made up public land (combined approx. 5,252 ha, 14%). METHODS Landscape Composition and Site Classification We analyzed composition of land cover types in all study townships in southwest and west-central Wisconsin using ArcGIS 9.2 (Environmental Systems Research Institute, Redlands, CA) and National Land Cover Data 2006 (NLCD; Fry et al. 2011). Land cover classes in the NLCD dataset included open water, developed area (low, medium, high intensity, and open space), barren land, deciduous forest, evergreen forest, mixed forest, shrub/scrub, grassland/herbaceous, hay/pasture, cultivated crops, woody wetlands, and emergent herbaceous wetlands. We aggregat- ed deciduous forest, mixed forest, evergreen forest, and woody wetland into a single forest cover class; likewise we combined grassland/herbaceous areas, hay/pasture, and cultivated crops into a single open cover class. We used Hawth’s Analysis Tools 3.27 (www.spatialecology.com/ htools, accessed 13 Sep 2010) and ArcGIS 9.2 to tabulate the area of all land cover classes. We chose to classify study townships dependent upon the dominant, aggregated land cover class within each township. Townships with > 50% forest cover were classified as forested study sites, and those with > 50% open cover were classified as open study sites. Therefore, we categorized Fairchild (70.4% forest, 18.7% open) and Stark (60.4% forest, 31.0% open) townships as forested sites, and Hale (32.1% forest, 59.5% open) and Westford (31.8% forest, 62.9% open) townships as open sites. Capture and Monitoring We trapped turkeys from January to mid-March, 2010 and 2011 using rocket nets (Bailey et al. 1980). We baited multiple sites within each study township with cracked corn and whole oats, and targeted female turkeys at each bait site. We determined age-class as adult or yearling from the shape and coloration of the outermost 2 primaries (Pelham and Dickson 1992), weighed individuals, and fitted all captured turkeys with individually numbered aluminum non-reward butt-end leg bands (National Band and Tag, Newport, KY). Prior to release, we also fitted females with backpack style radio transmitters with 8-hour mortality switches (Model A1540, Advanced Telemetry Systems, Isanti, MN) attached with 3.8-mm shock cord (Recreational Equipment, Inc., Madison, WI). Transmitters weighed 80 g and had an expected battery life of 2,172 days. All birds were handled and released at the capture site in accordance with the University of Wisconsin-Madison Research Animal Care and Use Committee (Protocols A01036 and A01464). We located radio-marked female turkeys with portable receivers (Model R2000, Advanced Telemetry Systems) and 3-element folding Yagi antennas. We obtained locations via triangulation and visual observations 3 times per week after 986 The Journal of Wildlife Management 78(6) 19372817, 2014, 6, Downloaded from https://wildlife.onlinelibrary.wiley.com/doi/10.1002/jwmg.749 by University Of Florida, Wiley Online Library on [27/08/2026]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License capture through August, and checked location and mortality status of each female every 3–4 weeks from September through February. We used Program RADIO- TRACKER (J. Cary, University of Wisconsin-Madison, personal communication) programmed within Microsoft Visual Basic 1 software (Microsoft, Redmond, WA) and the maximum likelihood estimation method (Lenth 1981) to plot triangulated locations. We calculated locations in the field from 3 bearings taken within 30 minutes of the first azimuth. We calculated error ellipses for each triangulation and censored locations with error ellipse areas of > 5 ha; after censoring a location, we immediately obtained new bearings. Estimation of Demographic Parameters Female survival and cause of mortality.— We used annual and seasonal survival intervals defined by Wright et al. (1996) to investigate potential landscape effects on end point survival rates of female turkeys. Survival intervals roughly corresponded to biological events in the annual cycle of female turkeys. We defined the biological year as 15 March to 14 March the following year. Spring (15 Mar–18 Jul) included movements from wintering grounds (which we termed dispersal), nesting, and early brood-rearing. Sum- mer-fall (19 Jul–21 Nov) comprised late nesting activities and brood-rearing, and winter (22 Nov–14 Mar) included the period of persistent snow cover. We located and Figure 1. Locations and landscape composition of Fairchild (Eau Claire County), Hale (Trempealeau County), Stark (Vernon County), and Westford (Richland County) study townships where efforts to capture eastern wild turkey females were concentrated in southwest and west-central Wisconsin, USA, 2010–2011. Pollentier et al. Wild Turkey Ecology in Contrasting Landscapes 987 19372817, 2014, 6, Downloaded from https://wildlife.onlinelibrary.wiley.com/doi/10.1002/jwmg.749 by University Of Florida, Wiley Online Library on [27/08/2026]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License recovered transmitters when the transmitter’s mortality switch was activated. We attempted to determine cause of death by using field sign, evidence from field necropsies, and marks left on recovered transmitters (Hygnstrom et al. 1994, Thogmartin and Schaeffer 2000). We classified cause of death as predation (avian, canid, or other mammal), harvest, other (e.g., accidental, disease, vehicle collision, weather- related), or unknown (carcass severely scavenged or inconclusive evidence). Because we could not reliably distinguish the difference between coyote ( Canis latrans ), red fox ( Vulpes vulpes ), gray fox ( Urocyon cinereoargenteus ), and domestic dog kills, we grouped predations deemed to be caused by these species into a canid category. Nesting and brood survival.— We suspected females to be laying or incubating when triangulation results suggested localization in the same area on 3 successive occasions. We determined nesting location by homing to suspected radio- marked females to verify nesting, determine clutch size, and record nest location. To minimize disturbance, we continued to monitor nesting females 3 times weekly at distances 100 m by triangulation, and assumed a nest was still active if locations remained constant. We determined nest fate when the female was no longer attending the nest. We classified a nest as successful by presence of hatched eggshells, or as failed if nest contents were depredated, destroyed, or abandoned. We evaluated depredated nests for evidence of predator species by examining the nest site and characteristics of eggshell fragments (Hygnstrom et al. 1994). We determined nest initiation date by backdating and examining telemetry data (Paisley et al. 1998). We conducted flush counts at approximately 2 and 4 weeks post-hatch to estimate poult survival, which have been shown to yield similar results as survival estimates from radio-marked turkey poults (Speake et al. 1985, Hubbard et al. 1999 b ). Statistical Analyses Female survival.— We used known fate models in Program MARK 6.1 (White and Burnham 1999) and the Kaplan– Meier product limit estimator modified for staggered entry (Pollock et al. 1989 a ) to estimate female survival at each study site. We used a 1-week time step starting 15 March and ending 14 March the following year to estimate annual and seasonal survival. We assumed that survival rates were independent among individuals, the censoring mechanism was random, and that newly marked female turkeys had the same survival function as previously radio-marked turkeys. We used left- and right-censoring mechanisms (Pollock et al. 1989 b ) to account for unknown fates due to slipped transmitters, transmitter failures, and survival to the following study year. The beginning annual sample size during the second field season included female turkeys surviving from the first year; we reclassified yearlings surviving to their second year as adults. Additionally, we attributed mortalities that occurred within 14 days of capture to be caused by capture-related effects and censored these individuals from survival analyses. We used an information- theoretic approach and based model selection and inference on minimization of Akaike’s Information Criterion adjusted for small sample size (AIC c ) and AIC c weights ( w i ) to evaluate the influence of study site, female age-class, season, and study year on weekly survival of female turkeys (Burnham and Anderson 2002). We considered models with D AIC c 2 to show substantial support (Burnham and Anderson 2002) and considered parameters in the best- supported models to be informative if the 95% confidence intervals for the b values excluded 0. We used model- averaging for parameter estimation to account for uncertain- ty in model selection (Burnham and Anderson 2002). We pooled data from seasons across both years of study and conducted independent survival analyses for townships classified as forest or open as described above. We used the most parsimonious model to derive seasonal survival probabilities, and estimated annual survival as the product of consecutive spring (15 Mar–18 Jul), summer-fall (19 Jul– 21 Nov), and winter (22 Nov–14 Mar) seasonal survival estimates. Productivity and poult survival.— We considered 8 April the beginning of the nesting season because this was the earliest nest-initiation date of a radio-marked female turkey during this study. We calculated nesting rate as the proportion of radio-marked females that attempted at least 1 nest, and calculated renesting rates as the percentage of radio-marked females that initiated a second nest conditional upon loss of the first nest. Apparent nesting and renesting rates are likely biased low because nests may fail before discovery. Therefore, we estimated the number of nests and renests using estimates of daily survival rates for each landscape type (forest or open), female age-class, and nesting attempt using a Horvitz–Thompson estimator (Dinsmore et al. 2002). We defined nest survival as the proportion of nests that produced 1 hatched poult, and we used the nest survival model in Program MARK 6.1 (White and Burnham 1999) to generate maximum likelihood estimates of daily nest survival corrected for exposure before discovery. We evaluated the influence of female age-class, landscape (forest or open), nesting attempt (first or renest), and year on nest survival, and used model selection and inference based on minimization of AIC c values (Burnham and Anderson 2002). We considered all models with D AIC c 2 to show substantial support, while D AIC c 4 showed some support (Burnham and Anderson 2002). We used model- averaging to account for model uncertainty. We used the most parsimonious model to estimate daily nest survival probabilities, and extrapolated the overall nest survival probability as the product of daily survival probabilities over a 40-day exposure period (12-day laying period plus 28-day incubation period). We calculated variances of extrapolated nest survival probabilities using the delta method (Seber 1982). We defined apparent nesting success as the proportion of female turkeys alive on 8 April that hatched a successful nest, even if it was a renest. We included females that were considered to have not attempted to nest in our nesting success analysis. Our definition of apparent nesting success is similar to the term hen success used elsewhere (e.g., Paisley et al. 1998) and takes into account the separate components of the nesting process (e.g., nesting rates and 988 The Journal of Wildlife Management 78(6) 19372817, 2014, 6, Downloaded from https://wildlife.onlinelibrary.wiley.com/doi/10.1002/jwmg.749 by University Of Florida, Wiley Online Library on [27/08/2026]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License nest survival) and combines them into a single estimate of reproductive success. We excluded females disturbed by observers during initial nesting attempts from renesting rate calculations (if the first nest was abandoned), nest survival (attempts were treated separately), and apparent nesting success calculations if the female did not attempt to renest. We defined hatching rate as the proportion of eggs that hatched in successful nests. Because hatching rate of eggs within a clutch likely is not independent, we estimated standard errors by the ratio method of Cochran (1963:29– 33). We used analysis of variance (ANOVA) to indepen- dently test for year and landscape (forest and open) differences in average clutch size and hatching rates. Additionally we tested for differences in frequency of birds nesting and apparent nesting success among female age- classes, years, and forested and open landscapes with 2-way contingency tables. We calculated interval survival estimates for poults observed during flush-count surveys. We based the initial sample size in each brood on the number of poults likely to have hatched and left the nest. We estimated survival from 0 to 2 weeks and 2 to 4 weeks post-hatch with a modified Kaplan–Meier estimator (Flint et al. 1995) that does not require independent survival among brood members. We used a bootstrap sampling procedure with 2,000 replicates to estimate standard errors associated with survival estimates (Flint et al. 1995). We excluded data if a radio-marked female was lost or died before we conducted a flush count. We pooled data across years and sites to compare survival rates between landscapes with the normal approximation ( Z ) according to Pollock et al. (1989 a ). We set significance thresholds at a ¼ 0.05 for all statistical tests. RESULTS We captured 138 female turkeys (40 yearlings and 98 ad) at the 4 study sites during 2010 and 2011. We fitted all but 9 females ( n ¼ 41, 31, 27, and 30 at Fairchild, Hale, Stark, and Westford townships, respectively) with radio trans- mitters to monitor survival and productivity from capture until death or until transmitters were lost or failed. We did not consider 15 birds because cause of death was attributed to capture-related effects within 14 days after initial marking. We right censored 4 additional females because we lost radio signal ( n ¼ 1) or birds slipped transmitters ( n ¼ 3). We monitored 58 female turkeys in 2010 and 75 females in 2011 for reproductive estimates, of which 80 nests were located and monitored during our study. Survival Seasonal and annual survival.— Seven of 20 yearling hens (35.0%) survived through the 2010–2011 field season and were reclassified as adult hens for the second field season, and 28 of 44 (63.6%) adult hens survived through the 2010–2011 field season. Seasonal variation in survival and landscape effects were important components within both competing models for female turkey survival and accounted for > 98% of the relative support (Table 1). The top model ( w 1 ¼ 0.61) also included an age-class effect; however, the beta estimate did not differ significantly from 0 ( b age ¼ 0.47, 95% CI: 0.05, 0.99). Yearling female survival was lower than that of adult females across all seasons (Table 2), and extrapolated annual survival across all study townships was 0.404 (95% CI: 0.229, 0.578) for yearlings and 0.607 (95% CI: 0.529, 0.685) for adults. A model excluding age-class effect was parsimonious ( D AIC c ¼ 1.00, w 2 ¼ 0.37; Table 1), and model parameter estimates of landscape effects between forested and open sites ( b landscape ¼ 0.99, 95% CI: 1.48, 0.49) and seasonal effects ( b season ¼ 0.77, 95% CI: 0.94, 0.59) were both informative. Seasonal survival was lower during the spring (15 Mar–18 Jul) nesting and early brood- rearing period than during the remainder of the year in both forested and open landscapes (Table 2). Model-averaged estimates of female turkey annual survival across all study townships was 0.515 0.035, and was lower in forested landscapes (2010 ¼ 0.450 0.075, 2011 ¼ 0.279 0.094) compared to open landscapes (2010 ¼ 0.707 0.053, 2011 ¼ 0.651 0.045; Fig. 2). Cause of mortality.— We recorded 72 deaths from March 2010 to March 2012 (Table 3), and attributed the majority of mortalities to predators (77.8%). We estimated canids caused at least 52 of 56 (92.9%) predator-related losses; we found 2 carcasses that were cached under downed trees, which we attributed to bobcats ( Lynx rufus ). Great- horned owls ( Bubo virginianus ) appeared to have killed 2 other radio-marked females. To compare the relative importance of predation, we pooled all non-predation losses into a single category. We found that cause of mortality did not differ by year ( x 2 1 ¼ 0 : 20, P ¼ 0.651); however, with years combined, cause of mortality did vary by season ( x 2 1 ¼ 9 : 82, P ¼ 0.007). Predators caused most mortalities Table 1. Candidate models and model statistics for weekly survival of adult and yearling eastern wild turkey females ( n ¼ 114) at 4 study townships in southwest and west-central Wisconsin, USA, 15 March 2010– 14 March 2012. Model structure a Model statistics b K AIC c D AIC c w i Dev. S season þ landscape þ female age 4 693.40 0.00 0.613 685.39 S season þ landscape 3 694.40 1.00 0.372 688.40 S season 2 701.80 8.40 0.015 697.80 S season þ female age 3 702.57 9.17 0.010 696.57 S year þ landscape þ female age 4 747.02 53.62 0.000 739.01 S female age þ landscape 3 748.77 55.37 0.000 742.77 S year þ landscape 3 749.35 55.95 0.000 743.35 S landscape 2 750.48 57.08 0.000 746.47 S year þ township 5 761.65 68.25 0.000 755.64 S township 4 762.52 69.12 0.000 758.51 S female age 2 763.27 69.87 0.000 759.27 S year 2 763.88 70.48 0.000 759.88 S constant 1 764.94 71.54 0.000 762.94 S week 52 771.30 77.60 0.000 666.25 a Season ¼ spring (15 Mar–18 Jul), summer-fall (19 Jul–21 Nov), or winter (22 Nov–14 Mar); landscape ¼ forest or open; female age ¼ adult or yearling; township ¼ Fairchild, Hale, Stark, or Westford; year ¼ 2010– 2011 or 2011–2012; week ¼ week of the year, constant ¼ constant survival. b Model fit is described by the number of parameters ( K ), Akaike’s Information Criterion corrected for small sample size (AIC c ), AIC c weights ( w i ), and deviance (Dev.). Pollentier et al. Wild Turkey Ecology in Contrasting Landscapes 989 19372817, 2014, 6, Downloaded from https://wildlife.onlinelibrary.wiley.com/doi/10.1002/jwmg.749 by University Of Florida, Wiley Online Library on [27/08/2026]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License during spring (87.5%), and fewer during summer-fall (69.2%) and winter (45.5%). Cause of predation also varied between the 2 types of landscapes ( x 2 1 ¼ 4 : 86, P ¼ 0.027); we estimated predators caused 85.4% of mortalities in forested landscapes, compared to 62.5% of mortalities in open landscapes (Table 3). Predators killed 7 female turkeys that were known to be actively incubating, all of which occurred in forested landscapes. Additionally, 6 females were killed with broods 2 weeks old, 5 in forested landscapes and 1 in an open landscape. Productivity Nesting chronology.— First-known nest initiation dates ranged from 9 April to 13 June ( n ¼ 29) during 2010 and 8 April to 3 June ( n ¼ 40) during 2011. The median date of initiation of first nests was 5 May ( n ¼ 34) for females in forested landscapes and 16 May ( n ¼ 35) for females in open landscapes. Renesting females initiated nests from 4 May to 7 June ( n ¼ 5) in 2010 and 12 May to 20 June ( n ¼ 8) in 2011. Hatching dates ranged from 21 May to 21 July ( n ¼ 13) in 2010 and 25 May to 21 July ( n ¼ 17) in 2011. Nesting rate and nesting success.— Nesting rates were higher for adults than yearlings ( x 2 1 ¼ 42 : 21, P 0.001); however, with female age-classes pooled we found little annual variation ( x 2 1 ¼ 3 : 43, P ¼ 0.064) or difference between landscapes ( x 2 1 ¼ 2 : 29, P ¼ 0.130; Table 4). Fourteen of 34 (41.2%) adult females attempted to renest following failure of a first nest; however, we found no yearlings attempted to renest. Adult renesting rates varied little annually ( x 2 1 ¼ 0 : 13, P ¼ 0.717) and were similar between landscapes ( x 2 1 ¼ 0 : 08, P ¼ 0.774). We modeled nest survival for the 105-day nesting season (8 Apr–21 Jul). The top model suggested that nest survival varied between years and had 33% of the relative support (Table 5). However, we found model uncertainty; a constant model ( D AIC c ¼ 1.32) and a model containing landscape effect ( D AIC c ¼ 1.36) were also parsimonious. Models where nest survival differed between nesting attempts and female age-classes showed some support ( D AIC c 4, w i < 0.09; Table 5). Model- averaged estimates of overall nest survival varied little between forested ( ^ u ¼ 0 : 273, SE ¼ 0.08) and open land- scapes ( ^ u ¼ 0 : 201, SE ¼ 0.07; Table 4). With female age- classes pooled, we detected little difference in apparent nesting success between years ( x 2 1 ¼ 0 : 62, P ¼ 0.431) or between landscapes ( x 2 1 ¼ 0 : 32, P ¼ 0.573; Table 4). Clutch size and hatching rate.— Average clutch size ( SE) of first nests was 10.67 0.33 eggs and was similar between adults and yearlings ( F 1,49 ¼ 0.004, P ¼ 0.948). Average Table 2. Seasonal survival estimates ð ^ S Þ of radio-marked adult and yearling (Yrl) eastern wild turkey females in southwest and west-central Wisconsin, USA, 15 March 2010–14 March 2012. Group a Season c No. individuals added b Spring Summer-fall Winter From 2010 to 2011 Ad Yrl n d ^ S SE n ^ S SE n ^ S SE 2010–2011 Forest 23 9 16 0.557 0.087 14 0.871 0.085 42 0.928 0.049 Open 21 11 28 0.875 0.058 27 0.964 0.036 42 0.838 0.066 2011–2012 Forest 13 19 10 22 0.525 0.076 16 0.719 0.097 12 0.740 0.111 Open 22 14 6 30 0.754 0.067 26 0.863 0.064 26 1.000 0.000 Age e Yrl 0.602 0.084 0.813 0.100 0.825 0.083 Ad 0.728 0.045 0.906 0.041 0.921 0.033 Means 0.672 0.039 0.861 0.036 0.890 0.031 a We pooled study sites according to our general land cover classifications, where Fairchild and Stark Townships were categorized as forested landscapes and Hale and Westford Townships were categorized as open landscapes. b Females alive on 15 March. Yearlings surviving through their second winter were reclassified as adults in 2011–2012. c We defined biological seasons as spring (15 Mar–18 Jul), summer-fall (19 Jul–21 Nov), and winter (22 Nov–14 Mar). d Sample size at the conclusion of the period. e Female age-classes were pooled across study sites and years. Figure 2. Annual survival rates (15 Mar–14 Mar) of radio-marked female eastern wild turkeys found in townships categorized as forested or open landscapes in southwest and west-central Wisconsin, USA, 2010–2012. Forested landscape townships are Fairchild and Stark Townships, and open landscape townships are Hale and Westford Townships. Beginning annual sample sizes on 15 March were n ¼ 62 for forest 2010–2011, n ¼ 42 for forest 2011–2012, n ¼ 52 for open 2010–2011, and n ¼ 42 for open 2011–2012. We omitted confidence intervals and standard errors for clarity. Vertical lines denote the change in biological seasons, from spring to summer-fall (19 Jul) and summer-fall to winter (22 Nov). 990 The Journal of Wildlife Management 78(6) 19372817, 2014, 6, Downloaded from https://wildlife.onlinelibrary.wiley.com/doi/10.1002/jwmg.749 by University Of Florida, Wiley Online Library on [27/08/2026]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License clutch size of second (9.82 0.46) and third nest attempts (10.50 1.50) were also similar to first nests ( F 2,59 ¼ 0.73, P ¼ 0.485). We found little difference in average clutch size among all nests across study sites ( F 3,58 ¼ 1.53, P ¼ 0.216). We did not include 1 suspected dump nest from Fairchild Township (forested landscape), which included 23 eggs in clutch size calculations. Among all study sites, the proportion of eggs laid in successful nests ( n ¼ 30) that survived incubation was high with hatching rates surpassing 0.92 (SE ¼ 0.03). Cause of nest failure.— Of 82 known nesting attempts, we recorded 52 (63.4%) nest losses. Mammalian predation was the main factor affecting nest success and accounted for 57.7% of all nest failures. Common mammalian nest predators in the study area included raccoon ( Procyon lotor ), striped skunk ( Mephitis mephitis ), Virginia opossum ( Didel- phis virginiana ), gray fox, red fox, and coyote. Other causes of nest failure included avian depredation (presumably Ameri- can crow [ Corvus brachyrhynchos ]; n ¼ 3, 5.8%), farming operation ( n ¼ 1, 1.9%), flooding ( n ¼ 1, 1.9%), abandon- ment ( n ¼ 7, 13.5%), unknown cause ( n ¼ 6, 11.5%), and disturbance by researchers ( n ¼ 4, 7.7%). When we pooled all non-predator- and predator-related losses into a single category, respectively, and found no difference in the cause of nest loss between forest and open landscapes ( x 2 1 ¼ 0 : 746, P ¼ 0.388). Table 3. Probable causes and number of eastern wild turkey female mortalities in southwest and west-central Wisconsin, USA, 15 March 2010–14 March 2012. Cause No. of mortalities Forest a Open Subtotal Spring b Summer-fall Winter Spring Summer-fall Winter Predator Canid 29 6 4 10 2 1 52 Bobcat 2 0 0 0 0 0 2 Avian 0 0 0 1 1 0 2 Disease 1 0 0 1 1 0 3 Accident c 2 0 0 2 0 0 4 Harvest d 0 2 1 0 0 0 3 Weather 0 0 1 0 0 0 1 Unknown 0 0 0 0 1 4 5 Total 34 8 6 14 5 5 72 a General land cover characteristics of study townships, where Fairchild and Stark Townships represent forested landscapes ( > 50% forest cover) and Hale and Westford Townships are open landscapes ( > 50% open cover). b Seasons were defined as spring (15 Mar–18 Jul), summer-fall (19 Jul–21 Nov), and winter (22 Nov–14 Mar). c Accidents were attributed to farming operations ( n ¼ 3) in hay fields during the nesting season and 1 lost to a vehicle collision. d Includes legally harvested females ( n ¼ 2) and 1 harvested outside the hunting season. Table 4. Reproductive parameter estimates for adult and yearling (Yrl) radio-marked eastern wild turkey females in southwest and west-central Wisconsin, USA, 2010–2011. Parameter a Nesting rate b Nest survival c Nesting success d First nest Renest First nest Renest n ^ u SE n ^ u SE n ^ u SE n ^ u SE n ^ u SE Age-specific Ad 101 0.901 0.03 34 0.412 0.09 60 0.228 0.06 13 0.362 0.16 97 0.330 0.05 Yrl 32 0.344 0.09 4 0.000 0.00 5 0.121 0.13 0 0.000 0.00 32 0.031 0.03 Year 2010 58 0.690 0.06 15 0.333 0.13 26 0.310 0.10 5 0.688 0.26 55 0.291 0.06 2011 75 0.827 0.04 23 0.319 0.10 39 0.160 0.06 8 0.174 0.14 74 0.230 0.05 Landscape Forest 63 0.825 0.05 19 0.368 0.11 32 0.250 0.08 7 0.393 0.21 61 0.279 0.06 Open 70 0.714 0.05 19 0.368 0.11 33 0.188 0.07 6 0.280 0.21 68 0.235 0.05 Overall 133 0.767 0.04 38 0.368 0.08 65 0.217 0.05 13 0.338 0.15 129 0.256 0.04 a Age-classes pooled across years and landscape categories. Year (2010 and 2011) is pooled across age-classes and landscapes. Landscape categories represent general land cover characteristics of study townships, where Fairchild and Stark Townships are forested landscapes ( > 50% forest cover) and Hale and Westford Townships are open landscapes ( > 50% open). b Proportion of females alive on 8 April that attempted to nest or renest, adjusted with Horvitz–Thompson estimators (Dinsmore et al. 2002) to account for nests that may have failed before discovery. c Nest survival estimates derived as the product of daily survival probabilities over a 40-day exposure period (12-day laying period plus 28-day incubation period). d Apparent nesting success, defined as the proportion of females alive on 8 April that successfully hatched a nest, even if it was a renest, and includes females that were considered to have not attempted a nest. Pollentier et al. Wild Turkey Ecology in Contrasting Landscapes 991 19372817, 2014, 6, Downloaded from https://wildlife.onlinelibrary.wiley.com/doi/10.1002/jwmg.749 by University Of Florida, Wiley Online Library on [27/08/2026]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License Poult survival.— During 2010–2011, we attempted to collect poult-count data from 30 radio-marked female turkeys that successfully hatched a nest. However, we excluded 6 broods from our analysis because the radio- marked female was lost or died before an initial flush count could be performed. We estimated survival with 224 poults in the remaining 24 broods. Most poult mortalities (85.2%, n ¼ 121) occurred during the 0- to 2-week interval post- hatch (Table 6). We acquired additional poult-counts at approximately 4 weeks post-hatch for 22 broods that remained. Estimated poult survival rates ( SE) at approxi- mately 4 weeks post-hatch were 0.379 0.111 ( n ¼ 116) in 2010 and 0.352 0.063 ( n ¼ 108) in 2011. Pooled across both years of study, survival was lower at 4 weeks post-hatch in forested landscapes compared to open landscapes ( Z ¼ 2.67, P ¼ 0.008; Table 6). DISCUSSION We compared vital rates of turkeys in open-agricultural and forest-dominated landscapes in southwest and west-central Wisconsin at the township level to remove any potential effects due to dissimilar spatial scale. Differences in survival estimates were evident between landscapes that were closely related to adult survival rates during the spring (15 Mar–18 Jul) nesting and early brood-rearing period. Overall reproductive success was fairly equal among all 4 study sites and across landscapes. Yearling nesting rates ( < 0.35) and apparent nesting success ( < 0.10) were both much lower than that of adult females and appeared to contribute little to the population. Our results indicate that conservation actions designed to maximize benefits for turkeys and turkey management in Wisconsin require that managers must be cognizant that large-scale patterns of forest and opening on landscapes have a large influence on turkey survival and should incorporate this idea when drafting wild turkey habitat management plans. Survival and Probable Causes of Mortality Although age-class was included in our top-ranked model of female survival, the effect was marginal because the 95% confidence intervals of yearling and adult survival overlapped. The age-specific difference in survival may have been stronger, but our sample was skewed towards adults, and the Table 5. Candidate models and model statistics for daily nest survival ( n ¼ 76 nests) of eastern wild turkey females in forested and open landscapes in southwest and west-central Wisconsin, USA, 2010–2011. Model structure a Model statistics b K AIC c D AIC c w i Dev. S year 2 285.18 0.00 0.330 281.17 S constant 1 286.50 1.32 0.170 284.50 S landscape þ year 3 286.54 1.36 0.167 280.52 S nest attempt 2 287.85 2.67 0.087 283.84 S female age 2 287.93 2.75 0.084 283.92 S landscape þ year þ female age 4 287.96 2.78 0.082 279.93 S landscape 2 288.02 2.83 0.080 284.01 a Year ¼ 2010 or 2011; female age ¼ adult or yearling; landscape ¼ forest or o