Spatio-temporal variation in male white-tailed deer harvest rates in Pennsylvania: Implications for estimating abundance

The performance of 2 popular methods that use age-at-harvest data to estimate abundance of white-tailed deer is contingent on assumptions about variation in estimates of subadult (1.5 yr old) and adult (≥2.5 yr old) male harvest rates. Auxiliary data (e.g., estimates of survival or harvest rates from radiocollared animals) can be used to relax some assumptions, but unless these population parameters exhibit limited temporal or spatial variation, these auxiliary data may not improve accuracy. Unfortunately maintaining sufficient sample sizes of radiocollared deer for parameter estimation in every wildlife management unit (WMU) is not feasible for most state agencies. We monitored the fates of 397 subadult and 225 adult male white-tailed deer across 4 WMUs from 2002 to 2008 using radio telemetry. We investigated spatial and temporal variation in harvest rates and investigated covariates related to the patterns observed. We found that most variation in harvest rates was explained spatially and that adult harvest rates (0.36–0.69) were more variable among study areas than subadult harvest rates (0.26–0.42). We found that hunter effort during the archery and firearms season best explained variation in harvest rates of adult males among WMUs, whereas hunter effort during only the firearms season best explained harvest rates for subadult males. From a population estimation perspective, it is advantageous that most variation was spatial and explained by a readily obtained covariate (hunter effort). However, harvest rates may vary if hunting regulations or hunter behavior change, requiring additional field studies to obtain accurate estimates of harvest rates. © 2011 The Wildlife Society.     

Influence of habitat features and hunter behavior on white-tailed deer harvest

Sport hunting may help in controlling cervid populations over large areas. As with natural predators, several environmental factors can influence sport harvest. A better understanding of the environmental variables that limit the efficiency of sport hunting could provide guidelines for more efficient wildlife management using hunting. We studied white-tailed deer (Odocoileus virginianus) hunting on a high deer density island where hunting was the sole form of predation. Our objective was to study the behavior of sport hunters and determine the habitat characteristics (e.g., abundance of deer forage, visibility of the deer from the hunter's point of view, and accessibility of the territory to hunters) that are associated with a successful harvest. We collected movements and harvest site location data from 477 hunters equipped with handheld Global Positioning System (GPS) units. Harvest sites were visited and characterized, along with a paired random site, to determine the environmental conditions associated with a successful hunt. We also developed a model to predict the daily number of deer seen by hunters considering weather conditions, hunter characteristics (e.g., age, experience), and date of hunting. We used the mean number of deer seen per hunter per day as a relative index of local density in each hunted territory. At both the site and landscape scales, the combination of visibility and access had a positive effect on the distribution of harvested deer. Habitat types with less visual obstruction from vegetation enabled hunters to see more deer in a given day. At the site scale, harvested deer were located in areas with a lower density of access routes compared to areas where hunters travelled throughout the day. Using an innovative approach of studying hunter behavior with GPS technology, digital maps, and questionnaires, we highlighted the factors associated with hunter success. Our study suggests that habitat characteristics could be modified to increase harvest by improving accessibility and visibility near roads. Creating openings in mature and regenerating forest near access roads could make sport hunting a more efficient management tool, but the potential impact of increased forage availability in forest openings should not be overlooked. © 2012 The Wildlife Society.     

Land use and dispersal influence mortality in white-tailed deer

Restoring male age structure in white-tailed deer populations has become an important objective for many state agencies aimed at improving herd dynamics. Limiting mortality in the yearling (1–2 yr old) age class is a primary consideration, and regional differences in climate, habitat characteristics, hunting regulations, and hunter behavior complicate the understanding of how specific factors influence the risk of mortality. We used Cox proportional hazard modeling to determine the effects of body size, mean distance to road, dispersal behaviors, use of forested land, and use of land open to public hunting on the risk of mortality for a population of radio-collared, yearling males (n = 76) in Sussex County, Delaware, USA. Annual survival averaged 0.55 (95% CI = 0.45–0.68), with harvest accounting for 79% (26/33) of all mortalities. Measurements of body size (chest girth, shoulder height, and total length; cm) influenced dispersal probability but not dispersal distance. The best approximating model for mortality risk included a covariate for landownership, whereby mortality risk increased on public land. Among males who dispersed, longer-distance dispersal was associated with reduced mortality, which contradicts previous research describing dispersal as a high-risk behavior. The effect of landownership on mortality risk has not been previously identified, especially when regulations regarding harvest of yearling males are similar between landownership types. We observed annual survival rates of 0.69 (95% CI = 0.57–0.82) for deer apparently using private land exclusively during the hunting season, and 0.20 (95% CI = 0.11–0.48) for deer that used public land during the hunting season. Survival rates on private land were comparable to those of other regions actively managing male age structure. These results suggest survival of yearling males in the region is influenced by hunter harvest and the risks associated with dispersal may be minimal in areas where harvest pressure is low, although hunter harvest on public land may limit male age structure on a localized scale. © The Wildlife Society, 2019     

Phenotypic and reproductive variation in female white-tailed deer: The role of harvest and environment

Animal density and stochastic environmental events affect physical development and recruitment of cervids. Biologists who manage cervid populations based on a density-dependent paradigm may need to also consider environmental effects. We analyzed 12- to 25-year time series of 3 harvested white-tailed deer populations in Mississippi to determine the relative influence of harvest and environmental factors on female reproduction and phenotypic quality. Using simple and multiple linear regression, we related body mass of 1.5-, 2.5-, and ≥3.5-year females, and 2.5- and ≥3.5-year percent lactation to variables representing deer harvest, growing season precipitation and temperatures, high-quality agronomic plantings, and flooding events. Response across populations varied greatly, with harvest variables explaining most, some, or no variation in body mass and percent lactation. Biologists should consider the potential influence of environmental factors on phenotypic and reproductive variation when making management decisions. © 2012 The Wildlife Society.     

Effect of male age structure on reproduction in white-tailed deer

Selective harvest regimes that create female-biased sex ratios can potentially lead to delayed breeding, reduced breeding synchrony, reduced productivity, and a female-biased sex ratio of offspring. These resulting changes in breeding behavior and population dynamics have potential to adversely affect population growth. In 2002, Pennsylvania implemented harvest regulation changes that reduced deer density (increased harvest of antlerless deer) and increased the number and age of antlered deer (implemented antler point restriction regulations) that resulted in a less female-biased sex ratio. We monitored date of conception, productivity (embryos/female), and sex ratio of embryos during 1999–2006 to test if timing of breeding occurred earlier and with greater synchrony, if productivity of females increased, and if the sex ratio of offspring would shift towards more males. Deer density decreased 23% and the adult (≥1.5 yr old) sex ratio declined from 2.30 to 1.95 females/male. The ratio of ≥2.5-year-old to 1.5-year-old males shifted towards more older males (1:3.7 in 2002 to 1:1.59 in 2006) and the ≥2.5-year-old male population increased from 41,853 during 1999–2001 to 54,064 by 2006. We found no evidence of any change in the timing or variability of date of conception, productivity, or offspring sex ratio. We conclude that harvest regulation changes implemented in Pennsylvania, USA, were insufficient to affect timing of breeding or population dynamics and that efforts by managers to identify a desired sex ratio or manipulate sex ratios to achieve management goals on a statewide scale will be challenging. © 2019 The Wildlife Society.     

Incorporating harvest rates into the sex-age-kill model for white-tailed deer

Although monitoring population trends is an essential component of game species management, wildlife managers rarely have complete counts of abundance. Often, they rely on population models to monitor population trends. As imperfect representations of real-world populations, models must be rigorously evaluated to be applied appropriately. Previous research has evaluated population models for white-tailed deer (Odocoileus virginianus); however, the precision and reliability of these models when tested against empirical measures of variability and bias largely is untested. We were able to statistically evaluate the Pennsylvania sex-age-kill (PASAK) population model using realistic error measured using data from 1,131 radiocollared white-tailed deer in Pennsylvania from 2002 to 2008. We used these data and harvest data (number killed, age-sex structure, etc.) to estimate precision of abundance estimates, identify the most efficient harvest data collection with respect to precision of parameter estimates, and evaluate PASAK model robustness to violation of assumptions. Median coefficient of variation (CV) estimates by Wildlife Management Unit, 13.2% in the most recent year, were slightly above benchmarks recommended for managing game species populations. Doubling reporting rates by hunters or doubling the number of deer checked by personnel in the field reduced median CVs to recommended levels. The PASAK model was robust to errors in estimates for adult male harvest rates but was sensitive to errors in subadult male harvest rates, especially in populations with lower harvest rates. In particular, an error in subadult (1.5-yr-old) male harvest rates resulted in the opposite error in subadult male, adult female, and juvenile population estimates. Also, evidence of a greater harvest probability for subadult female deer when compared with adult (≥2.5-yr-old) female deer resulted in a 9.5% underestimate of the population using the PASAK model. Because obtaining appropriate sample sizes, by management unit, to estimate harvest rate parameters each year may be too expensive, assumptions of constant annual harvest rates may be necessary. However, if changes in harvest regulations or hunter behavior influence subadult male harvest rates, the PASAK model could provide an unreliable index to population changes. © 2012 The Wildlife Society.     

Evaluating the effect of predators on white-tailed deer: Movement and diet of coyotes

Coyotes (Canis latrans) may affect adult and neonate white-tailed deer (Odocoileus virginianus) survival and have been implicated as a contributor to the decline of deer populations. Additionally, coyote diet composition is influenced by prey availability, season, and region. Because coyote movement and diet vary by region, local data are important to understand coyote population dynamics and their impact on prey species. In southeast Minnesota, we investigated the effect of coyotes on white-tailed deer populations by documenting movement rates, distances moved, and habitats searched by coyotes during fawning and nonfawning periods. Additionally, we determined survival, cause-specific mortality, and seasonal diet composition of coyotes. From 2001 to 2003, we captured and radiocollared 30 coyotes. Per-hour rate of movement averaged 0.87 km and was greater (P = 0.046) during the fawning (1.07 km) than the nonfawning period (0.80 km); areas searched were similar (P = 0.175) between seasons. Coyote habitat use differed during both seasons; habitats were not used in proportion to their availability (P < 0.001). Croplands were used more (P < 0.001) than their proportional availability during both seasons. Use of grasslands was greater during the fawning period (P = 0.030), whereas use of cropland was greater in the nonfawning period (P < 0.001). We collected 66 fecal samples during the nonfawning period; coyote diets were primarily composed of Microtus spp. (65.2%), and consumption of deer was 9.1%. During the study, 19 coyotes died; annual survival rate range was 0.33–0.41, which was low compared with other studies. Consumption of deer was low and coyotes searched open areas (i.e., cropland) more than fawning areas with dense cover. These factors in addition to high coyote mortality suggested that coyote predation was not likely limiting white-tailed deer populations in southeast Minnesota. © 2011 The Wildlife Society.     

Contribution of dogs to white-tailed deer hunting success

Dogs (Canis familiaris) are used in hunting white-tailed deer (Odocoileus virginianus) in 10 North American jurisdictions. Although the practice is longstanding and controversial, the effects of dogs on the outcome of the hunt have rarely been studied. We evaluated the influence of dogs on recreational hunting of white-tailed deer based on long-term data from southeastern Ontario, Canada. Over 25 years, annual surveys of hunters were used to collect data on hunting effort and deer harvest from approximately 85 camps, roughly half of which had dogs. We investigated the relationship between harvest and 3 treatments (i.e., 0 dogs, 1 dog, and ≥2 dogs in camp), interactions with weather and deer density, and effects of neighboring camps. Dogs enhanced hunter success. We found no difference in deer encounter rates but, per unit effort, camps with ≥2 dogs harvested 0.013 (26%) more deer per hunter-day, missed 0.010 (23%) more deer per hunter-day, and wounded 0.002 (40%) more deer per hunter-day than camps without dogs. Conversely, camps without dogs saw, without shooting at, 0.033 (23%) more deer per hunter-day than camps with ≥2 dogs. These results are consistent with the idea that hunters with dogs are less selective. Hunters with dogs harvested more fawns per unit effort, but we found no difference in the harvest rate of older female deer. More precipitation, greater wind speed, lower temperatures and greater deer density improved harvest success but had no differential effect among dog treatments. Hunter success at camps with ≥2 dogs was less when neighboring camps also had ≥2 dogs. Because antlerless deer quotas are the principal means to control populations, increasing use of hunting dogs is unlikely to have substantial effects in managing overabundant deer. © 2012 The Wildlife Society.     

Using spotlight observations to predict resource selection and abundance for white-tailed deer

Spotlight surveys for white-tailed deer (Odocoileus virginianus) can yield large presence-only datasets applicable to a variety of resource selection modeling procedures. By understanding how populations distribute according to a given resource for a reference area, density and abundance can be predicted across new areas assuming the relationship between habitat quality (measured by an index of selection) and species distribution are equivalent. Habitat-based density estimators have been applied to wildlife species and are useful for addressing conservation and management concerns. Although achieving reliable population estimates is a primary goal for spotlighting studies, presence-only models have yet to be applied to spotlight data for estimating habitat selection and abundance for deer. From 2012 to 2017, we conducted spring spotlight surveys in each of 99 counties in Iowa, USA, and collected spatial locations for 20,149 groups of deer (n = 71,323 individuals). We used a resource selection function (RSF) based on deer locations to predict the relative probability of use for deer at the population level and to estimate statewide abundance. The number of deer observed statewide increased significantly with increasing RSF value for all years and the mean RSF value along survey transects explained 59% of the variability in county-level deer counts, indicating that a functional response between habitat quality and deer distribution existed at landscape scales. We applied our RSF to a habitat-based density estimator (extrapolation) and zero-inflated Poisson (ZIP) and negative binomial (ZINB) count models to predict statewide abundance from spotlight counts. Population estimates for 2012 were variable, indicating that atypical weather conditions may affect spotlight counts and population estimates in some years. For 2013–2017, we predicted a mean population of 439,129 (95% CI ∼ ± 55,926), 440,360 (∼ ± 43,676), and 465,959 (∼ ± 51,242) deer across years for extrapolation, ZIP, and ZINB models, respectively. Estimates from all models were not significantly different than estimates from an existing deer population accounting model in Iowa for 2013 and 2016, and differed by <76,000 deer for all models from 2013–2017. Extrapolation and ZIP models performed similarly and differed by <2,897 deer across all years, whereas ZINB models showed inconsistencies in model convergence and precision of estimates. Our results indicate that presence-only models are capable of producing reliable and precise estimates of resource selection and abundance for deer at broad landscape scales in Iowa and provide a tool for estimating deer abundance in a spatially explicit manner. © 2019 The Wildlife Society.     

Spatial distribution and landscape associations of large-antlered deer

For more than a century, the Boone and Crockett Club has kept records of harvest locations for 38 categories of North American big game, which includes measurement details and sizes of enduring characteristics (e.g., antlers, skulls, horns). All white-tailed deer (Odocoileus virginianus) entered in the Club records (i.e., record deer) have large antlers because the measurements must meet or exceed a minimum size to qualify for entry. We used the records from the Club to test the hypothesis that the spatial distribution in the harvest of record deer was related to the number of antlered deer harvested, variations in land productivity, soil mineral content, and attributes of land cover for 9 states in the midwestern United States. We used a Bayesian theoretical framework to develop a spatial model that examined the influence of explanatory variables on the number of record deer harvested in each county. In our study area, 3,658 record deer were harvested in 692 of the 856 counties (80.8%). More record deer were harvested in counties that had more high-contrast edges, less contiguous land cover, and greater variation in soil productivity. These results provide information for managers and hunters to better understand the spatial distribution of record deer and factors that are correlated with their distribution. © 2019 The Wildlife Society.     

Genetic diversity and population structure in urban white-tailed deer

Habitat fragmentation in urban areas has left many species isolated and vulnerable to loss of genetic variation. White-tailed deer (Odocoileus virginianus), however, thrive in urban areas. We compared genetic diversity and structure among deer in 2 urban metroparks with deer in a fenced reserve and with deer from an open, continuously distributed population to inform urban deer management. If urban deer maintain species' typical matrilineal genetic structure, removal of female groups may effectively reduce local abundance. However, if gene flow in urban areas is high, dispersal may impede efforts to reduce abundance. Although genetic diversity was high and mean relatedness was near zero in all locations, distributions of pairwise relatedness in urban metroparks and the fenced reserve contained greater proportions of closely and distantly related deer than the open locations, likely attributable to matrilineal structure. In addition, deer from the metroparks (approx. 65 km apart) were moderately differentiated (F_st = 0.092) indicating gene flow in urban areas may be less than in other landscapes. Our results indicate that removal of matrilineal groups may reduce local urban deer abundance without inducing immigration from surrounding areas. © The Wildlife Society, 2012     

Genetic consequences of white-tailed deer (Odocoileus virginianus) restoration in Mississippi

White-tailed deer (Odocoileus virginianus) were nearly extirpated from the southeastern USA during the late 19th and early 20th centuries. Recovery programmes, including protection of remnant native stocks and transplants from other parts of the species' range, were initiated in the early 1900's. The recovery programmes were highly successful and deer are presently numerous and continuously distributed throughout the southeastern USA. However, the impact of the recovery programmes on the present genetic structure of white-tailed deer remains to be thoroughly investigated. We used 17 microsatellite DNA loci to assess genetic differentiation and diversity for 543 white-tailed deer representing 16 populations in Mississippi and three extra-state reference populations. There was significant genetic differentiation among all populations and the majority of genetic variation (> or = 93%) was contained within populations. Patterns of genetic structure, genetic similarity and isolation by distance within Mississippi were not concordant with geographical proximity of populations or subspecies delineations. We detected evidence of past genetic bottlenecks in nine of the 19 populations examined. However, despite experiencing genetic bottlenecks or founder events, allelic diversity and heterozygosity were uniformly high in all populations. These exceeded reported values for other cervid species that experienced similar population declines within the past century. The recovery programme was successful in that deer were restored to their former range while maintaining high and uniform genetic variability. Our results seem to confirm the importance of rapid population expansion and habitat continuity in retaining genetic variation in restored populations. However, the use of diverse transplant stocks and the varied demographic histories of populations resulted in fine-scale genetic structuring.     

Evaluating how hunters see and react to telemetry collars on white-tailed deer

Fates of individuals outfitted with radiotransmitters commonly are used for estimating survival rates in populations of large animals that are hunted. Despite precautions, this practice may be subject to complex biases associated with hunter reaction to presence of radiotransmitters. To assess this potential bias we conducted an experiment using artificial deer (i.e., decoys) to measure hunters' abilities to see deer and determine if deer seen were wearing radiocollars. We used logistic regression to quantify probabilities that seeing deer and subsequently seeing radiocollars might be influenced by distance, percent visual obstruction, body orientation, hunter experience, and antler characteristics of deer. Additionally, we evaluated how experience and antler characteristics of deer might influence a hunter's decision to harvest a radiocollared deer. We found that 25.8% of the potentially observable collared deer (n = 663) were subsequently observed by hunters. Odds of observing deer and radiocollars increased 95% and 230%, respectively, for each additional log(yr) of hunting experience. Willingness to harvest radiocollared deer increased 89% for each additional log(yr) of hunting experience and 144% for large-antlered deer relative to antlerless deer. When hunting is an important source of mortality, analysts need to understand how potential biases associated with observing deer are associated with hunters' reactions to and subsequent decisions to harvest radiocollared animals. Our study suggested that presence of radiocollars may influence a deer's potential risk of being harvested and in turn bias telemetry-based estimates of survival, given that hunting mortality is the largest component of total mortality in hunted deer populations. Collar-based telemetry is used nearly universally by wildlife managers and researchers throughout North America and elsewhere to estimate and monitor the survival of big game populations that are managed through hunting. Our findings demonstrate that these estimates are likely subject to complex and systematic biases that managers should consider when evaluating future population-level effects of managed hunting. © 2011 The Wildlife Society     

Survival of white-tailed deer fawns on Marine Corps Base Quantico

Some jurisdictions in the eastern United States have reduced harvest of white-tailed deer (Odocoileus virginianus) because of perceived declines in recruitment and population size over the last decade. Although the restoration of American black bears (Ursus americanus) and the colonization of coyotes (Canis latrans) have increased fawn predation in some areas, limited information exists on how temporally dynamic resources and weather influence fawn survival. Therefore, we evaluated fawn survival probability, cause specific mortality, and if factors such as oak (Quercus spp.) mast abundance, winter severity, precipitation, and landscape composition influenced mortality risk on Marine Corps Base Quantico in northern Virginia, USA, from 2008 to 2019. We tracked 248 fawns outfitted with very high frequency radio-collars and predation was the leading cause of mortality (n = 42; 45%). We estimated survival to 133 days and survival pooling all years (2008–2019) was 0.50 (95% CI = 0.42–0.60). Increased annual red oak (Quercus spp.) mast abundance from the previous fall reduced mortality hazard for fawns. The longevity of our study revealed a link between fawn survival and a specific maternal resource (red oak mast) only available during gestation. Our results highlight the importance of oak mast in eastern deciduous forests and, more broadly, overwinter maternal condition on white-tailed deer recruitment.