A fine-scale examination of parturition timing in temperate ungulates.

Parturition timing has long been a topic of interest in ungulate research. However, few studies have examined parturition timing at fine scale (e.g., <1 day). Predator activity and environmental conditions can vary considerably with diel timing, which may result in selective pressure for parturition to occur during diel times that maximize the likelihood of neonate survival. We monitored parturition events and early-life survival of elk (Cervus canadensis) and mule deer (Odocoileus hemionus) in Utah, USA to better understand diel timing of parturition in temperate ungulates. Diel timing of parturition was moderately synchronous among conspecifics and influenced by environmental variables on the date of parturition. For elk, parturition events were most common during the morning crepuscular period and generally occurred later (i.e., closer to 12:00) when a relatively large proportion of the moon was illuminated. For mule deer, parturition events were most common during the diurnal period and generally occurred later (i.e., closer to 15:00) on cold, wet dates. Diel timing of parturition did not influence neonate survival, but larger datasets may be required to verify the apparent lack of influence. Although additional work could evaluate alternative variables that might affect parturition timing, our data provide an improved and finer scale understanding of reproductive ecology and phenology in ungulates.

Evaluating movement-based methods for estimating the frequency and timing of parturition in mule deer.

BACKGROUND: Information on reproduction of harvested species such as mule deer (Odocoileus hemionus) is vital for conservation and management. Furthermore, parturition in ungulates may be detected using patterns of movement logged by GPS transmitters. Several movement-based methods have been developed to detect parturition in ungulates including the Peterson method, behavioral change point analysis (BCPA), rolling minimum convex polygons (rMCP), individual-based method (IBM), and population-based method (PBM). Our objectives were to (1) test the accuracy and the precision of each previously described method and (2) develop an improved method optimized for mule deer that incorporated aspects of the other methods. METHODS: We determined parturition timing and status for female mule deer fitted with GPS collars and implanted with vaginal implant transmitters (VITs). We used movement patterns before and after parturition to set movement thresholds for each movement-based method. Following model training, we used location and birth date data from an external dataset to test the effectiveness of each movement-based method. Additionally, we developed a novel method for detecting parturition called the analysis of parturition indicators (API). We used two regression analyses to determine the accuracy and precision of estimates generated by each method. RESULTS: The six methods we employed varied in accuracy, with the API, rMCP, and BCPA being most accurate. Precision also varied among methods, with the API, rMCP, and PBM generating the most precise estimates of parturition dates. The API and the rMCP performed similarly and better overall than any of the other existing methods. CONCLUSIONS: We found that movement-based methods could be used to accurately and precisely detect parturition in mule deer. Further, we determined that the API and rMCP methods had the greatest overall success at detecting parturition in mule deer. The relative success of the API and rMCP may be attributed to the fact that both methods use home range size to detect parturition and are validated using known parturition dates of collared deer. We present the API as an efficient method of estimating birth status and timing of parturition of mule deer fitted with GPS transmitters, as well as affirm the effectiveness of a previously developed method, rMCP.

Estimating age of mule deer in the field: Can we move beyond broad age categories?

Age of individuals is an intrinsic demographic parameter used in the modeling and management of wildlife. Although analysis of cementum annuli from teeth is currently the most accurate method used to age ungulates, the age of live ungulates in the field can be estimated by examining tooth wear and tooth replacement patterns. However, there may be limitations to aging based on tooth wear as the rate of tooth wear likely varies among individuals due to factors such as age, diet, environment, and sex. Our objective was to determine the reliability of estimating age for mule deer based on tooth wear and tooth replacement patterns. We compared ages estimated by tooth wear (collected at time of capture for a statewide monitoring effort) to ages determined from cementum analysis (from teeth collected after mortalities of radio-tracked animals from the monitoring effort). Accuracy was high; ages estimated from tooth wear were within one year of cementum ages >75% of the time when aged by experienced observers. Bias in accuracy for estimates of age was low but slightly biased toward underestimation (i.e., 0.6 years on average)-especially as cementum age increased. Our results indicate that aging mule deer using patterns in tooth wear can be reliable if observers estimating age have experience using this method.

Bighorn sheep show similar in-host responses to the same pathogen strain in two contrasting environments.

Ecological context-the biotic and abiotic environment, along with its influence on population mixing dynamics and individual susceptibility-is thought to have major bearing on epidemic outcomes. However, direct comparisons of wildlife disease events in contrasting ecological contexts are often confounded by concurrent differences in host genetics, exposure histories, or pathogen strains. Here, we compare disease dynamics of a Mycoplasma ovipneumoniae spillover event that affected bighorn sheep populations in two contrasting ecological contexts. One event occurred on the herd's home range near the Rio Grande Gorge in New Mexico, while the other occurred in a captive facility at Hardware Ranch in Utah. While data collection regimens varied, general patterns of antibody signal strength and symptom emergence were conserved between the two sites. Symptoms appeared in the captive setting an average of 12.9 days postexposure, average time to seroconversion was 24.9 days, and clinical signs peaked at approximately 36 days postinfection. These patterns were consistent with serological testing and subsequent declines in symptom intensity in the free-ranging herd. At the captive site, older animals exhibited more severe declines in body condition and loin thickness, higher symptom burdens, and slower antibody response to the pathogen than younger animals. Younger animals were more likely than older animals to clear infection by the time of sampling at both sites. The patterns presented here suggest that environment may not be a major determinant of epidemiological outcomes in the bighorn sheep-M. ovipneumoniae system, elevating the possibility that host- or pathogen-factors may be responsible for observed variation.

Variation in movement patterns of mule deer: have we oversimplified migration?

BACKGROUND: Conservation and management of migratory animals has gained attention in recent years, but the majority of research has focused on stereotypical 'migrant' and 'resident' behaviors, often failing to incorporate any atypical behaviors or characterize migratory behaviors beyond distance and timing of the migration. With migration threatened by anthropogenic development and climate change, it is crucial that we understand the full range of migratory behaviors. Our objective was to demonstrate and characterize the variation in migration strategies, including typical and atypical migratory behaviors for mule deer (Odocoileus hemionus) in Utah, USA. METHODS: Because calculation of common metrics such as distance, timing, and use of stopovers during migration did not adequately describe the variation we observed in migratory behavior for this species-particularly when animals visited multiple (> 3) ranges for extended lengths of time-we developed additional methods and categories to describe observed variation in migratory behavior. We first categorized trajectories based on the number of discrete, separate ranges and range shifts between them. Then, we further characterized the variation in migration strategies by examining the timing, duration, and distance traveled within each of the categories. We also examined if and how frequently individual deer switched among categories from year to year. RESULTS: We classified 1218 movement trajectories from 722 adult female mule deer, and found that 54.4% were dual-range migrants, who made one round-trip to one distinct range. Multi-range migrants (23.6%) made one round-trip during which they stayed at multiple discrete ranges. Commuters (1.0%) traveled to the same range multiple times, and poly migrants (1.5%) made multiple round-trips to different ranges. Gradual movers (2.5%) did not show a discrete range shift but moved gradually between ranges, whereas residents (12.6%) never left their home ranges, and dispersers (4.4%) left but never returned. Of the deer that we monitored for multiple years, 51.2% switched among categories. CONCLUSION: We conclude that the substantial number of atypical migratory strategies, as well as the number of deer that switched categories, underlines the importance of studying these less-stereotyped behaviors that may be exhibited by large proportions of populations. Acknowledging and investigating the full complexity and diversity in migratory strategies might uncover unknowns with respect to underlying factors and drivers of migration, and can help shape effective conservation strategies.

The influence of habitat use on harvest vulnerability of cow elk (Cervus canadensis).

Pressure from hunting can alter the behavior and habitat selection of game species. During hunting periods, cervids such as elk (Cervus canadensis) typically select for areas further from roads and closer to tree cover, while altering the timing of their daily activities to avoid hunters. Our objective was to determine the habitat characteristics most influential in predicting harvest risk of elk. We captured 373 female elk between January 2015 and March 2017 in the Uinta-Wasatch-Cache National Forest and surrounding area of central Utah, USA. We determined habitat selection during the hunting season using a resource selection function (RSF) for 255 adult cow elk. Additionally, we used a generalized linear mixed model to evaluate risk of harvest based on habitat use within home ranges (3rd order selection) as well as the location of the home range on the landscape to evaluate vulnerability on a broader scale. Female elk selected for areas that reduced hunter access (rugged terrain, within tree cover, on private land). Age, elevation and distance to roads within a home range were most influential in predicting harvest risk (top model accounted for 36.2% of AIC weight). Elevation and distance to trees were most influential in predicting risk when evaluating the location of the home range (top model accounted for 42.1% of AIC weight). Vulnerability to harvest was associated with proximity to roads. Additionally, survival in our landscape decreased with age of femaleelk.

Spatial processes decouple management from objectives in a heterogeneous landscape: predator control as a case study.

Predator control is often implemented with the intent of disrupting top-down regulation in sensitive prey populations. However, ambiguity surrounding the efficacy of predator management, as well as the strength of top-down effects of predators in general, is often exacerbated by the spatially implicit analytical approaches used in assessing data with explicit spatial structure. Here, we highlight the importance of considering spatial context in the case of a predator control study in south-central Utah. We assessed the spatial match between aerial removal risk in coyotes (Canis latrans) and mule deer (Odocoileus hemionus) resource selection during parturition using a spatially explicit, multi-level Bayesian model. With our model, we were able to evaluate spatial congruence between management action (i.e., coyote removal) and objective (i.e., parturient deer site selection) at two distinct scales: the level of the management unit and the individual coyote removal. In the case of the former, our results indicated substantial spatial heterogeneity in expected congruence between removal risk and parturient deer site selection across large areas, and is a reflection of logistical constraints acting on the management strategy and differences in space use between the two species. At the level of the individual removal, we demonstrated that the potential management benefits of a removed coyote were highly variable across all individuals removed and in many cases, spatially distinct from parturient deer resource selection. Our methods and results provide a means of evaluating where we might anticipate an impact of predator control, while emphasizing the need to weight individual removals based on spatial proximity to management objectives in any assessment of large-scale predator control. Although we highlight the importance of spatial context in assessments of predator control strategy, we believe our methods are readily generalizable in any management or large-scale experimental framework where spatial context is likely an important driver of outcomes.

Large herbivores surf waves of green-up during spring.

The green wave hypothesis (GWH) states that migrating animals should track or 'surf' high-quality forage at the leading edge of spring green-up. To index such high-quality forage, recent work proposed the instantaneous rate of green-up (IRG), i.e. rate of change in the normalized difference vegetation index over time. Despite this important advancement, no study has tested the assumption that herbivores select habitat patches at peak IRG. We evaluated this assumption using step selection functions parametrized with movement data during the green-up period from two populations each of bighorn sheep, mule deer, elk, moose and bison, totalling 463 individuals monitored 1-3 years from 2004 to 2014. Accounting for variables that typically influence habitat selection for each species, we found seven of 10 populations selected patches exhibiting high IRG-supporting the GWH. Nonetheless, large herbivores selected for the leading edge, trailing edge and crest of the IRG wave, indicating that other mechanisms (e.g. ruminant physiology) or measurement error inherent with satellite data affect selection for IRG. Our evaluation indicates that IRG is a useful tool for linking herbivore movement with plant phenology, paving the way for significant advancements in understanding how animals track resource quality that varies both spatially and temporally.

Effects of climate and plant phenology on recruitment of moose at the southern extent of their range.

Climate plays a fundamental role in limiting the range of a species, is a key factor in the dynamics of large herbivores, and is thought to be involved in declines of moose populations in recent decades. We examined effects of climate and growing-season phenology on recruitment (8-9 months old) of young Shiras moose (Alces alces shirasi) over three decades, from 18 herds, across a large geographic area encompassing much of the southern extent of their range. Recruitment declined in 8 of 18 herds during 1980-2009, whereas others did not exhibit a temporal trend (none showed a positive trend). During those three decades, seasonal temperatures increased, spring-summer precipitation decreased, and spring occurred earlier, became shorter in duration, and green-up occurred faster. Recruitment was influenced negatively by warm temperatures during the year before young were born, but only for herds with declining recruitment. Dry spring-summers of the previous year and rapid rates of spring green-up in the year of birth had similar negative influences across declining and stable herds. Those patterns indicate both direct (year(t)) and delayed (year(t-1)) effects of weather and plant phenology on recruitment of young, which we hypothesize was mediated through effects on maternal nutritional condition. Suppressed nutrition could have been induced by (1) increased thermoregulatory costs associated with warming temperatures and (2) shortened duration of availability of high-quality forage in spring. Progressive reductions in net energetic gain for species that are sensitive to climate may continue to hamper individual fitness and population dynamics.