Revisions of Rump Fat and Body Scoring Indices for Deer,Elk, and Moose

Abstract: Because they do not require sacrificing animals, body condition scores (BCS), thickness of rump fat (MAXFAT), and other similar predictors of body fat have advanced estimating nutritional condition of ungulates and their use has proliferated in North America in the last decade. However, initial testing of these predictors was too limited to assess their reliability among diverse habitats, ecotypes, subspecies, and populations across the continent. With data collected from mule deer (Odocoileus hemionus), elk (Cervus elaphus), and moose (Alces alces) during initial model development and data collected subsequently from free-ranging mule deer and elk herds across much of the western United States, we evaluated reliability across a broader range of conditions than were initially available. First, to more rigorously test reliability of the MAXFAT index, we evaluated its robustness across the 3 species, using an allometric scaling function to adjust for differences in animal size. We then evaluated MAXFAT, rump body condition score (rBCS), rLIVINDEX (an arithmetic combination of MAXFAT and rBCS), and our new allometrically scaled rump-fat thickness index using data from 815 free-ranging female Roosevelt and Rocky Mountain elk (C. e. roosevelti and C. e. nelsoni) from 19 populations encompassing 4 geographic regions and 250 free-ranging female mule deer from 7 populations and 2 regions. We tested for effects of subspecies, geographic region, and captive versus free-ranging existence. Rump-fat thickness, when scaled allometrically with body mass, was related to ingesta-free body fat over a 38–522-kg range of body mass (r² = 0.87; P < 0.001), indicating the technique is remarkably robust among at least the 3 cervid species of our analysis. However, we found an underscoring bias with the rBCS for elk that had >12% body fat. This bias translated into a difference between subspecies, because Rocky Mountain elk tended to be fatter than Roosevelt elk in our sample. Effects of observer error with the rBCS also existed for mule deer with moderate to high levels of body fat, and deer body size significantly affected accuracy of the MAXFAT predictor. Our analyses confirm robustness of the rump-fat index for these 3 species but highlight the potential for bias due to differences in body size and to observer error with BCS scoring. We present alternative LIVINDEX equations where potential bias from rBCS and bias due to body size are eliminated or reduced. These modifications improve the accuracy of estimating body fat for projects intended to monitor nutritional status of herds or to evaluate nutrition's influence on population demographics.     

Genetic Diversity and Divergence among Bighorn Sheep from Reintroduced Herds in Washington and Idaho

Bighorn sheep (Ovis canadensis) populations in the western United States have undergone widespread declines and extirpations since the late nineteenth century as a consequence of introduced diseases, competition with livestock, and unregulated hunting. Washington, Idaho, USA, and British Columbia, Canada were historically thought to be occupied by 2 bighorn lineages or subspecies: Rocky Mountain (O. c. canadensis) and California (O. c. californiana). The putative California lineage was completely extirpated in the United States, and reintroductions to reestablish populations were sourced directly or indirectly from a single region in southern British Columbia. Restoration efforts have attempted to maintain the diversity and divergence of these 2 lineages, sometimes referred to as subspecies although taxonomic classifications have changed over time. In this study we describe genetic variation in a subset of native and reintroduced herds of California and Rocky Mountain bighorn sheep. We examined genetic diversity and divergence between bighorn sheep herds using 15 microsatellite loci, including 4 loci linked to genes involved in immune function. We analyzed 504 samples from reintroduced herds in Washington (n = 10 California herds, n = 4 Rocky Mountain herds) and Idaho (n = 5 California), and source herds in Oregon (n = 1 Rocky Mountain) and British Columbia (n = 5 California, 1 Rocky Mountain). Genetic structure reflected known reintroduction history, and geographic proximity also was associated with decreased genetic divergence. Herds in Washington and Idaho sourced from California bighorn sheep were less genetically diverse than those sourced from Rocky Mountain herds. Also, levels of relatedness within and across California herds were higher than in Rocky Mountain herds and similar to what would be expected for full and half siblings. Lower diversity and higher relatedness among California herds is a concern for long-term fitness and likely related to past population bottlenecks, fewer source populations, and management history, such as entirely sourcing California herds from British Columbia. Genetic divergence of neutral loci between California and Rocky Mountain herds was greater than that of adaptive loci, potentially indicating that balancing selection has maintained similar genetic diversity across lineages in loci associated with immune and other adaptive functions. Thus, we recommend future reintroductions and augmentations should continue to use source populations from the appropriate California or Rocky Mountain lineage to avoid potential outbreeding depression and maintain possible adaptive differences. This could be accomplished by obtaining sheep from ≥1 source within the genetic lineage, while avoiding sourcing from admixed herds. Future work encompassing a broader geographic sampling of populations and a greater portion of the genome is necessary to better evaluate the degree to which contemporary divergence between lineages is associated with recent founder effects and genetic isolation or evolutionary adaptation. © 2021 The Wildlife Society     

Characterization of 11 polymorphic tetranucleotide microsatellites for forensic applications in California elk (Cervus elaphus canadensis)

Eleven polymorphic tetranucleotide microsatellite loci have been developed for forensic use in the protection of California elk. Based on a reference sample of elk taken from three races throughout California, the loci consist of 4–9 alleles (average 6.125). Probabilities of identity ( Paetkau et al. 1995 ) range from 0.079 to 0.288, with an overall probability of identity of 1.3 × 10^?9 (one in 7.8 × 10⁸).     

Genetic Characteristics of Restored Elk Populations in Kentucky

Translocations are a common management practice to restore or augment populations. Understanding the genetic consequences of translocation efforts is important for the long-term health of restored populations. The restoration of elk (Cervus canadensis) to Kentucky, USA, included source stocks from 6 western states, which were released at 8 sites in southeastern Kentucky during 1997–2002. We assessed genetic diversity in restored herds and compared genetic similarity to source stocks based on 15 microsatellite DNA loci. Genetic variation in the restored populations was comparable to source stocks ( allelic richness = 3.52 and 3.50; expected heterozygosity = 0.665 and 0.661 for restored and source, respectively). Genetic differentiation among all source and restored populations ranged from 0.000 to 0.065 for pairwise F_ST and 0.034 to 0.161 for pairwise Nei's D_A. Pairwise genetic differentiation and Bayesian clustering revealed that stocks from Utah and North Dakota, USA, contributed most to restored populations. Other western stocks appeared less successful and were not detected with our data, though our sampling was not exhaustive. We also inferred natural movements of elk among release sites by the presence of multiple genetic stocks. The success of the elk restoration effort in Kentucky may be due, in part, to the large number of elk (n = 1,548), repeated releases, and use of diverse source stocks. Future restoration efforts for elk in the eastern United States should consider the use of multiple stock sources and a large number of individuals. In addition, preservation of genetic samples of founder stock will enable detailed monitoring in the future. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

The efficacy of highway wildlife collision mitigation in preventing elk mortality in central Ontario

In the Burwash area of north-central Ontario, Canada, expansion of the Trans-Canada highway from 2 to 4 lanes was accompanied by installation of a range of wildlife collision-mitigation infrastructure (e.g., exclusion fencing, underpasses). To assess the overall effectiveness of these measures, we monitored the spatial distribution and mortality rates of elk (Cervus canadensis) prior to and following highway expansion, distinguished by season (winter, snowfree) and corridor-type (highway, railway). We measured herd-level risk by the proportion of positions falling within 200-m railway and highway buffer zones using Bayesian methods. Spatial analysis confirmed that there was a distinct northward shift in the winter distribution of elk following construction, situating the elk past the north end of the exclusion fence. This increased the herd's exposure to highway traffic by 3.6 times (proportion of points before = 0.0041 ± 0.002 [SE], after = 0.0147 ± 0.003, P = 0.005), and resulted in a more than 2-fold increase in elk road mortality from 0.6 elk/yr/20 km during 8 years prior to implementation to 1.5 elk/yr/20 km during 8 years after implementation. Exposure to railways remained unchanged and consistently higher than highway exposure regardless of season (e.g., post-mitigation, winter proportion of points = 0.0453 ± 0.005), matched by consistently high mortality counts (proportion of points before = 6.4 elk/yr/20 km, after = 6.6 elk/yr/20 km). Our results demonstrate that while wildlife-vehicle collision mitigation is generally beneficial to wildlife and humans, failure to account for the local characteristics of wildlife populations can lead to suboptimal mitigation designs that reduce their effectiveness and lead to unintended wildlife impacts.     

Microsatellite analysis of North American wapiti (Cervus elaphus) populations

Eleven populations of wapiti (Cervus elaphus) were analysed for genetic diversity using 12 microsatellite loci. Samples were taken from Vancouver Island, British Columbia; Burwash and French River herds in Ontario; Ya Ha Tinda Ranch, Alberta; and Banff, Elk Island, Jasper, Kootenay, Riding Mountain, Yellowstone and Yoho National Parks. Overall, wapiti populations have on average three to four alleles per locus and an average expected heterozygosity that ranged from 25.75 to 52.85%. The greatest genetic distances were observed between the Vancouver population and all other populations. Using the assignment test, Roosevelt wapiti (C. e. roosevelti Merriam 1897) assigned only to the Vancouver Island population. The distance and assignment values suggest a divergence of the Roosevelt wapiti from other populations and support the subspecific status for the Vancouver Island population. No evidence was found for the existence of unique Eastern wapiti (C. e. canadensis Erxleben 1777) in the Burwash or French River herds in Ontario. The overlapping distribution of genotypes from indigenous populations from Riding Mountain, Elk Island and Yellowstone National Parks suggests that wapiti were once a continuous population before settlers decimated their numbers. The lack of differentiation between these populations raises questions about the status of Manitoban (C. e.manitobensis Millais 1915) and Rocky Mountain (C. e.nelsoni Bailey 1935) subspecies.     

Assessment of Elk Highway Permeability by Using Global Positioning System Telemetry

Abstract: Highways have significant direct and indirect impact on natural ecosystems, including wildlife barrier and fragmentation effects, resulting in diminished habitat connectivity and highway permeability. We used Global Positioning System (GPS) telemetry to assess Rocky Mountain elk (Cervus elaphus nelsoni) permeability across a 30-km stretch of highway in central Arizona, USA, currently being reconstructed with 11 wildlife underpasses, 6 bridges, and associated ungulate-proof fencing. The highway was reconstructed in phases, allowing for comparison of highway crossing and passage rates during various stages of reconstruction. We instrumented 33 elk (25 F, 8 M) with GPS receiver collars May 2002 to April 2004. Our collars accrued 101,506 GPS fixes with 45% occurring within 1 km of the highway. Nearly 2 times the proportion of fixes occurred within 1 km of the highway compared with random. We think elk were attracted to the highway corridor by riparian—meadow foraging habitats that were 7 times more concentrated within the 1-km zone around the highway compared with the mean proportion within elk use areas encompassing all GPS fixes. Elk crossed the highway 3,057 times; crossing frequency and distribution along the highway were aggregated compared with random. Crossing frequency within 0.16-km highway segments was negatively associated with the distance to riparian—meadow habitats (r_s = -0.714, n = 190, P < 0.001). Mean observed crossing frequency (92.6 ± 23.5 [SE] crossings/elk) was lower than random (149.6 ± 27.6 crossings/elk). Females crossed 4.5 times as frequently as males. Highway permeability among reconstruction classes was assessed using passage rates (ratio of highway crossings to approaches); our overall mean passage rate was 0.67 ± 0.08 crossings per approach. The mean passage rate for elk crossing the highway section where reconstruction was completed (0.43 ± 0.15 crossings/approach) was half that of sections under reconstruction and control sections combined (0.86 ± 0.09 crossings/approach). Permeability was jointly influenced by the size of the widened highway and associated vehicular traffic on all lanes. Crossing frequency was used to delineate where ungulate-proof fencing yielded maximum benefit in intercepting and funneling crossing elk toward underpasses, promoting highway safety. Use of passage rates provides a quantitative measure to assess permeability, conduct future pre- and postconstruction comparisons, and to develop mitigation strategies to minimize highway impacts to wildlife.     

The Influence of Translocation Strategy and Management Practices on the Genetic Variability of a Reestablished Elk (Cervus elaphus) Population

Reintroduction of terrestrial vertebrates with the goal of ecosystem restoration typically establishes small and isolated populations that may experience reduced genetic variability due to founder effects and genetic drift. Understanding the genetic structure of these populations and maintaining adequate genetic diversity is important for long‐term restoration success. We quantified genetic variability at six microsatellite loci for a reintroduced population of Cervus elaphus (elk) restored to the tallgrass prairie ecosystem of northeastern Kansas. Allelic richness, observed and expected heterozygosity were intermediate to levels reported in other North American elk populations. Current levels of genetic variability in restored North American elk populations were not well explained by founding population size, number of founding populations, or number of years since the last translocation. Simulation results suggest that the retention of genetic variability in isolated populations is strongly influenced by mating system while also being impacted by temporal variability in population size and population growth rate. Our results have implications for understanding how translocation strategies and post‐reintroduction management may influence genetic variability in restored populations.     

Influences on Release-Site Fidelity of Translocated Elk

Several eastern states are considering the restoration of free‐ranging elk populations via translocation from western populations. Optimal habitat immediately surrounding release sites has been found to enhance elk reintroduction success in western states. Little information exists, however, to aid eastern managers in identifying release sites with the highest chance of restoration success. We monitored the movements of 415 translocated elk released at three sites in southeastern Kentucky to identify landscape characteristics that enhance release‐site fidelity. The distance elk moved after release differed among sites (F_2,322 = 4.63, p = 0.01), age classes (F_2,322 = 4.37, p = 0.01), and time intervals (F_2,322 = 40.74, p < 0.001). At 6 and 12 months post‐release, adults (15.81 ± 17.32 and 16.38 ± 20.29) and yearlings (13.91 ± 16.44 and 14.61 ± 21.11) moved farther than calves (8.06 ± 14.03 and 9.37 ± 14.40). The release site with the highest fidelity was privately owned, 15% open, and had the highest amount of edge compared with the other release sites. The two remaining sites contained large amounts of expansive openland or forest cover with lower amounts of edge. Additionally, both sites were publicly owned and experienced a higher degree of human‐generated disturbance compared with the site to which elk were most faithful. When selecting release sites, managers should avoid areas dominated by a single cover type with little interspersion of other habitats. Rather, areas with high levels of open‐forest edge (approximately 5.0 km/km²) and limited‐human disturbance will likely enhance release‐site fidelity and promote restoration success.     

Hiding Without Cover? Defining Elk Security in a Beetle-Killed Forest

Recent mountain pine beetle (Dendroctonus ponderosae; pine beetle) outbreaks in the western United States have affected nearly 18 million ha of pine (Pinus spp.) forest and are unprecedented in spatial extent, severity, and duration, yet little is known about wildlife responses to large-scale insect outbreaks. Elk (Cervus canadensis) are important wildlife whose dominant management paradigm on public lands has focused on providing security habitat to increase survival during hunting seasons and to maintain elk presence on public lands to promote hunter opportunity. To assess the effect of pine beetles and associated changes in forest structure on elk security, we used a time series to characterize canopy cover pre- and post-pine beetle outbreak, characterized relative canopy cover among the dominant forest types in the study area post-pine beetle outbreak, and used global positioning system location data from male and female elk to define habitat relationships and security during the archery and rifle hunting seasons. Our study area was within the Elkhorn Mountains of southwest Montana, USA, 2015–2017, which experienced 80% mortality of lodgepole pine (Pinus contorta) forests during a pine beetle outbreak that peaked in 2008. We observed an 8.5% reduction in canopy cover within pine beetle-infested lodgepole pine forests, yet canopy cover remained relatively high among other forest types post-outbreak. The top-ranked habitat security models contained positive relationships with canopy cover, distance to motorized routes, terrain ruggedness, and slope with few notable differences among sexes and seasons. Across sexes and seasons, 75% and 50% of elk use was within areas with average canopy cover values ≥31 ± 6.65 (SD)% and ≥53 ± 5.7% that were an average of ≥2,072 ± 187.93 m and ≥3,496 ± 157.32 m from a motorized route, respectively. Therefore, we recommend fall elk security be defined as areas that meet these criteria for minimum canopy cover and distance from motorized routes in the Elkhorn Mountains and in other landscapes with similar forest characteristics and hunting pressures. Although we observed expected reductions in canopy cover within pine beetle-infested forests, defoliation alone did not appear to negatively affect elk security or reduce canopy cover below our management recommendations. Nonetheless, because of the prevalence of standing dead trees in our study area, we recommend future work that investigates the relationships with pine beetle-infested areas post-blowdown because changes in ground structure and costs of locomotion may affect elk habitat and security. © 2019 The Authors. Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Postrelease Dispersal of Reintroduced Elk (Cervus elaphus) in Ontario,Canada

We studied 2 years of postrelease telemetry data of elk (Cervus elaphus) translocated to their historic range limit in Ontario, Canada and sought to determine if postrelease movements were related to behavior, demography of released animals, or site–specific attributes such as length of holding period. During 1998–2004 we radio‐tracked 341 elk in 10 release groups via ground and aerial telemetry and monitored movement patterns relative to gender, age, and pre‐release holding period (4–112 days). We found that elk that were held for short periods prior to release (4–11 days) moved longer distances than those subject to extended conditioning (17–112 days), suggesting that an extended conditioning period is beneficial from the standpoint of promoting philopatry. When all elk were pooled by sex and age class, male calves remained in closer proximity (8.0 ± 13.2 km) to release sites than adult females (19.1 ± 20.6 km), adult males (19.7 ± 15.1 km), and female calves (14.4 ± 20.4 km). Most calves dispersed in a southeasterly direction whereas adults tended to travel southwest. Our results reveal that elk movement characteristics are influenced by factors such as release protocol and group demographics; these findings provide further insight regarding appropriate release methods for restoring natural populations near their historical range limit.     

Development of a Sightability Model for Low-Density Elk Populations in Ontario,Canada

ABSTRACT The status of recolonizing elk (Cervus elaphus) populations in Ontario, Canada, is unclear and there is a need for effective population survey methods that can be applied locally. We sought to develop a sightability model that could account for both low densities of elk and dense forest cover in elk-release areas in Ontario. We corrected winter aerial survey counts for sightability based on radiocollared animals known to be within observable distance of the aircraft. The multivariate model with the highest Akaike's Information Criterion corrected for sample size weight (w_i = 0.427) revealed that elk group size, elk activity, dominant tree type, percent canopy cover, and percent conifer cover were significant predictors of elk sightability. The group-size effect indicated that odds of sighting an elk increased by 1.353 (95% CI = 0.874-3.689) for every additional elk. Standing elk were 5.033 (95% CI = 0.936-15.541) times more likely to be observed than were resting elk, and those located in conifer cover were 0.013 (95% CI = 0.001-0.278) times less likely to be sighted than elk in deciduous cover. Furthermore, elk located in >50% canopy cover and >50% conifer cover were 0.041 (95% CI = 0.003-0.619) times and 0.484 (95% CI = 0.024-9.721) times less likely to be sighted than elk in more open habitat, respectively. During model validation, observers detected 79% (113/143) of known elk in any given area, and population and sightability model predictions (±90% CI) overlapped with the population estimate, implying that our predictive model was robust. Unsurprisingly, large groups of elk in open habitat increased model precision, which highlights difficulties of counting Ontario elk in their northern range. We conclude that our model provided increased reliability for estimating elk numbers in Ontario compared to existing methods, and that the estimator may be useful in other areas where elk density is low and sightability is poor due to dense forest cover.     

Influence of interspecific competition on mule deer birthing and rearing site selection

Ungulates often alter behavior and space use in response to interspecific competition. Despite observable changes in behavior caused by competitive interactions, research describing the effects of competition on survival or growth is lacking. We used spatial modeling to determine if habitat use by female mule deer (Odocoileus hemionus) was affected by other ungulate species prior to, during, and after parturition. We conducted our study in the Book Cliffs region of eastern Utah, USA, during 2019 and 2020. We used resource selection function (RSF) analysis to model space use of 4 ungulate species that potentially competed with mule deer: bison (Bos bison), cattle, elk (Cervus canadensis), and feral horses. We incorporated RSF models for competing species into a random forest analysis to determine if space use by mule deer was influenced by these other ungulate species. We used survival and growth data from neonate mule deer to directly assess potential negative effects of other ungulates. Habitat use by elk was an important variable in predicting use locations of mule deer during birthing and rearing. The relationship was positive, suggesting interference competition was not occurring. Survival of neonate mule deer increased as the probability of use by elk increased (hazard ratio = 0.185 ± 0.497 [SE]). Further, probability of use by elk in rearing habitat had no influence on growth of neonate mule deer from birth to 6 months of age, suggesting that exploitative competition was not occurring.     

Abiotic,bottom-up,and top-down influences on recruitment of Rocky Mountain elk in Oregon: A retrospective analysis

Understanding the relative effects of the many factors that may influence recruitment of ungulates is fundamental to managing their populations. Over the last 4 decades, average recruitment in some populations of elk (Cervus elaphus) in Oregon, USA declined from >50 to <20 juveniles per 100 females, and several competing hypotheses address these declines. We developed a priori models and constructed covariates spanning 1977–2005 from hunter-killed elk, elk population estimates, cougar harvest, and weather statistics to evaluate abiotic, bottom-up, and top-down factors that may explain annual variation and long-term trends of pregnancy, juveniles-at-heel in late autumn, and recruitment of juvenile elk in spring. In models of pregnancy status, August precipitation, age, and cougar index had positive effects, whereas previous year (t − 1) winter severity or winter precipitation_(t−1) and elk density had negative effects. In models of juvenile-at-heel in late autumn, August precipitation, August precipitation_(t−1), cougar index × elk density_(t−1), and age had positive effects, whereas cougar index, elk density_(t−1), and winter precipitation_(t−1) had negative effects. Juvenile recruitment was best explained by positive effects of August precipitation_(t−1), lactation rate, and cougar index × elk density_(t−1) and negative effects of cougar index and elk density_(t−1). Winter severity, precipitation, and temperature were not significant in explaining variation in elk recruitment. Annual variation in pregnancy, juvenile-at-heel, and recruitment was most influenced by August precipitation, whereas long-term trends in recruitment were most influenced by cougar densities with relatively weak effects of elk density. These results provide insight into causes of year-to-year and long-term trends of elk recruitment and provide a basis for more rigorous evaluation of factors affecting recruitment of elk. © 2012 The Wildlife Society.     

Habitat Selection by Female Elk During Minnesota's Agricultural Season

Understanding space-use patterns by elk (Cervus canadensis) is essential to alleviating human-elk conflicts, particularly when crop depredation by elk can make it harder to justify to the public the need to restore elk populations to regions with agricultural landscapes. In 2016–2017, we used global positioning system data from 20 female elk to investigate their selection for agricultural cover during the agricultural season (1 May–31 Oct) in northwestern Minnesota, USA. We estimated resource selection functions with resource availability defined at the home range scale. Elk space use was primarily determined by distance to forest cover in areas proximate to agricultural fields. During diurnal periods, elk selected areas with forest cover near agricultural fields planted with legumes and cereal. During nocturnal periods, elk selected for agricultural fields with little to no canopy and that were planted with legumes. We suggest that management of elk in northwestern Minnesota will require practices that discourage the use of agriculture by elk while improving natural habitats within areas managed for elk restoration. We suggest that forestry practices (i.e., thinning and burning) could improve cover and forage openings for elk in restoration areas. Furthermore, managers could work with agricultural producers where elk occur to plant crops favored by elk (i.e., legumes) outside known home ranges and plant fields within home ranges with crops that elk avoided (i.e., hay). Collectively, these practices may shift the ranges of elk herds into restoration areas and lower conflict between the public and elk restoration efforts. © 2020 The Wildlife Society.     

Habitat Selection of Rocky Mountain Elk in a Nonforested Environment

Abstract: Recent expansions by Rocky Mountain elk (Cervus elaphus) into nonforested habitats across the Intermountain West have required managers to reconsider the traditional paradigms of forage and cover as they relate to managing elk and their habitats. We examined seasonal habitat selection patterns of a hunted elk population in a nonforested high-desert region of southwestern Wyoming, USA. We used 35,246 global positioning system locations collected from 33 adult female elk to model probability of use as a function of 6 habitat variables: slope, aspect, elevation, habitat diversity, distance to shrub cover, and distance to road. We developed resource selection probability functions for individual elk, and then we averaged the coefficients to estimate population-level models for summer and winter periods. We used the population-level models to generate predictive maps by assigning pixels across the study area to 1 of 4 use categories (i.e., high, medium-high, medium-low, or low), based on quartiles of the predictions. Model coefficients and predictive maps indicated that elk selected for summer habitats characterized by higher elevations in areas of high vegetative diversity, close to shrub cover, northerly aspects, moderate slopes, and away from roads. Winter habitat selection patterns were similar, except elk shifted to areas with lower elevations and southerly aspects. We validated predictive maps by using 528 locations collected from an independent sample of radiomarked elk (n = 55) and calculating the proportion of locations that occurred in each of the 4 use categories. Together, the high- and medium-high use categories of the summer and winter predictive maps contained 92% and 74% of summer and winter elk locations, respectively. Our population-level models and associated predictive maps were successful in predicting winter and summer habitat use by elk in a nonforested environment. In the absence of forest cover, elk seemed to rely on a combination of shrubs, topography, and low human disturbance to meet their thermal and hiding cover requirements.     

The Restoration of Elk (Cervus elaphus) in Ontario, Canada: 1998–2005

In 1997, a plan to restore Elk (Cervus elaphus) to Ontario was approved by the provincial government. The objective of the Ontario elk restoration program, a multipartnered collaboration, was to restore a species that had been extirpated from the province during the 1800s. During 1998–2001, 460 elk were acquired from Elk Island National Park, Alberta, for release in four areas of Ontario. As greater than 90% of the elk were radio collared, monitoring provided detailed information on the dynamics of the four populations. Comprehensive research projects using graduate students were implemented to determine the environmental impact of releasing elk in Ontario. Those studies are in progress or have been completed and include the effect of wolf predation on restored elk, white‐tailed deer and elk resource overlap, the development of genetic profiles for elk, and solutions for elk/human conflicts. Mortality of the released elk averaged 41% (190/460) during 1998–2004 with annual mortality generally declining over time in each release area. The primary causes of elk mortality included wolf predation (25% of mortalities), illegal shooting (13%), stress‐related emaciation (13%) (partially due to the stress of relocation), bacterial infections (7%), and collisions with vehicles (6%). Productivity has been high in one of the release areas with 24–65% of the cows being observed with calves during late winter surveys. However, productivity has been low in two of the northern release areas due to a variety of factors including wolf predation. In some areas, dispersion of elk appeared to be related to the length of time animals were kept in pens prior to release. The precalving population estimate for Ontario in March 2004 was 375–440 elk. A comprehensive program review was conducted in 2003/2004 that included recommendations relating to the future management of elk in Ontario.     

Fertility Control in Free-Ranging Elk Using Gonadotropin-Releasing Hormone Agonist Leuprolide: Effects on Reproduction,Behavior, and Body Condition

ABSTRACT Overabundant elk (Cervus elaphus) populations have become a significant problem in many areas of North America. This is particularly true for protected areas where high densities of elk can cause long-term ecological degradation. When lethal control is not acceptable in these environments, resource managers must often consider alternative methods for reducing the size of resident elk populations. A potential management alternative is controlling the fertility of female elk. A promising new approach to wildlife contraception involves the use of biodegradable implants containing the gonadotropin-releasing hormone (GnRH) agonist leuprolide. During fall 2002-spring 2004, we compared pregnancy rates, reproductive behavior, daily activity patterns, and body condition of 17 free-ranging female elk treated with a leuprolide formulation with 17 untreated females, in Rocky Mountain National Park, Colorado, USA. After treatment, the pregnancy rate of treated elk was 0%, whereas 79% of control elk became pregnant. The effects of treatment were reversed the subsequent year with the pregnancy rate of treated females 100% compared with 91% for controls. Reproductive behaviors were similar for treated and control elk during the breeding and postbreeding seasons; treated elk did not exhibit postrut reproductive behaviors. Moreover, we found no differences in daily activity patterns of experimental groups during the breeding or postbreeding seasons. Treated elk did not show improved body condition over pregnant females. Instead, treated females catabolized proportionately more body fat during winter after treatment and at a higher rate than pregnant control elk. However, this effect was reversed the next spring with no difference in body fat between treated and control elk. We conclude from this experiment that leuprolide, administered as a controlled release formulation, offers a safe and effective method of controlling fertility in free-ranging female elk. However, practical application is limited by treatment duration and the need to treat females before the breeding season.     

Intervarietal and intravarietal genetic structure in Douglas‐fir: nuclear SSRs bring novel insights into past population demographic processes,phylogeography, and intervarietal hybridization

Douglas‐fir (Pseudotsuga menziesii) is one of numerous wide‐range forest tree species represented by subspecies/varieties, which hybridize in contact zones. This study examined the genetic structure of this North American conifer and its two hybridizing varieties, coastal and Rocky Mountain, at intervarietal and intravarietal level. The genetic structure was subsequently associated with the Pleistocene refugial history, postglacial migration and intervarietal hybridization/introgression. Thirty‐eight populations from the USA and Canada were genotyped for 13 nuclear SSRs and analyzed with simulations and traditional population genetic structuring methods. Eight genetic clusters were identified. The coastal clusters embodied five refugial populations originating from five distinct refugia. Four coastal refugial populations, three from California and one from western Canada, diverged during the Pleistocene (56.9–40.1 ka). The three Rocky Mountain clusters reflected distinct refugial populations of three glacial refugia. For Canada, ice covered during the Last Glacial Maximum, we present the following three findings. (1) One refugial population of each variety was revealed in the north of the distribution range. Additional research including paleodata is required to support and determine whether both northern populations originated from cryptic refugia situated south or north of the ice‐covered area. (2) An interplay between intravarietal gene flow of different refugial populations and intervarietal gene flow by hybridization and introgression was identified. (3) The Canadian hybrid zone displayed predominantly introgressants of the Rocky Mountain into the coastal variety. This study provides new insights into the complex Quaternary dynamics of this conifer essential for understanding its evolution (outside and inside the native range), adaptation to future climates and for forest management.     

Temporal Overlap Among Feral Horses,Cattle, and Native Ungulates at Water Sources

Feral horse (Equus ferus caballus) populations on public rangelands in the western United States threaten forage production for livestock and wildlife habitat. Interference competition between feral horses and heterospecifics at watering sources can have negative effects on livestock and wildlife. Researchers have documented altered timing and behavior of wild ungulates at water sources when horses were present. The few studies examining these interactions have infrequently occurred within areas specifically managed for feral equids and have not occurred in sites with cattle. We used motion-sensitive cameras at 8 watering sources to document watering activity patterns and construct indices of temporal overlap among feral horses, cattle, elk (Cervus canadensis), mule deer (Odocoileus hemionus), and pronghorn (Antilocapra americana) within the Adobe Town Herd Management Area in southern Wyoming, USA, between June and September 2018 and 2019. Feral horses, cattle, and pronghorn exhibited a high degree of temporal overlap (>79%) in water use, with feral horses and pronghorn exhibiting the highest estimated percent overlap (88.1%, 95% CI = 86.5–89.6%). Mule deer and elk watering activity also overlapped with horses and cattle but to a lesser degree (<55%). Feral horses spent a mean of 16.7 ± 30.5 (SD) minutes during a watering event and were present at a given water source on average 4.5 ± 6.3% and up to 34.9% of the day, which is less than reported in previous studies. Cattle spent on average 23.5 ± 44.9 minutes during a watering event, and were present on average 4.2 ± 7.7% and up to 42.4% of the day at a single water source. Results of generalized linear mixed-effects models indicated that number of conspecifics was the strongest predictor of visit duration for pronghorn and horses; hour of the day and group size of heterospecifics were informative, but less important, variables. There was no difference in peak visitation time for any species between sites of high versus low horse or cattle use. Despite temporal overlap, we did not find evidence of interference competition between feral horses, cattle, and pronghorn. We recommend future examination of interference competition and its biological consequences between introduced and native ungulates at water sources of varying size across sites, equid population levels, and livestock stocking rates. © 2020 The Wildlife Society.