Pathology of Chronic Mycoplasma ovipneumoniae Carriers in a Declining Bighorn Sheep (Ovis canadensis) Population.

Bighorn sheep (Ovis canadensis) across North America commonly experience population-limiting epizootics of respiratory disease. Although many cases of bighorn sheep pneumonia are polymicrobial, Mycoplasma ovipneumoniae is most frequently associated with all-age mortality events followed by years of low recruitment. Chronic carriage of M. ovipneumoniae by adult females serves as a source of exposure of naïve juveniles; relatively few ewes may be responsible for maintenance of infection within a herd. Test-and-remove strategies focused on removal of adult females with evidence of persistent or intermittent shedding (hereafter chronic carriers) may reduce prevalence and mitigate mortality. Postmortem confirmation of pneumonia in chronic carriers has been inadequately reported and the pathology has not been thoroughly characterized, limiting our understanding of important processes shaping the epidemiology of pneumonia in bighorn sheep. Here we document postmortem findings and characterize the lesions of seven ewes removed from a declining bighorn sheep population in Wyoming, USA, following at least two antemortem detections of M. ovipneumoniae within a 14-mo period. We confirmed that 6/7 (85.7%) had variable degrees of chronic pneumonia. Mycoplasma ovipneumoniae was detected in the lung of 4/7 (57.1%) animals postmortem. Four (57.1%) had paranasal sinus masses, all of which were classified as inflammatory, hyperplastic lesions. Pasteurella multocida was detected in all seven (100%) animals, while Trueperella pyogenes was detected in 5/7 (71.4%). Our findings indicate that not all chronic carriers have pneumonia, nor do all have detectable M. ovipneumoniae in the lung. Further, paranasal sinus masses are a common but inconsistent finding, and whether sinus lesions predispose to persistence or result from chronic carriage remains unclear. Our findings indicate that disease is variable in chronic M. ovipneumoniae carriers, underscoring the need for further efforts to characterize pathologic processes and underlying mechanisms in this system to inform management.

Spatial prey availability and pulsed reproductive tactics: Encounter risk in a canid-ungulate system.

Predation risk is a function of spatiotemporal overlap between predator and prey, as well as behavioural responses during encounters. Dynamic factors (e.g. group size, prey availability and animal movement or state) affect risk, but rarely are integrated in risk assessments. Our work targets a system where predation risk is fundamentally linked to temporal patterns in prey abundance and behaviour. For neonatal ungulate prey, risk is defined within a short temporal window during which the pulse in parturition, increasing movement capacity with age and antipredation tactics have the potential to mediate risk. In our coyote-mule deer (Canis latrans-Odocoileus hemionus) system, leveraging GPS data collected from both predator and prey, we tested expectations of shared enemy and reproductive risk hypotheses. We asked two questions regarding risk: (A) How does primary and alternative prey habitat, predator and prey activity, and reproductive tactics (e.g. birth synchrony and maternal defence) influence the vulnerability of a neonate encountering a predator? (B) How do the same factors affect behaviour by predators relative to the time before and after an encounter? Despite increased selection for mule deer and intensified search behaviour by coyotes during the peak in mule deer parturition, mule deer were afforded protection from predation via predator swamping, experiencing reduced per-capita encounter risk when most neonates were born. Mule deer occupying rabbit habitat (Sylvilagus spp.; coyote's primary prey) experienced the greatest risk of encounter but the availability of rabbit habitat did not affect predator behaviour during encounters. Encounter risk increased in areas with greater availability of mule deer habitat: coyotes shifted their behaviour relative to deer habitat, and the pulse in mule deer parturition and movement of neonatal deer during encounters elicited increased speed and tortuosity by coyotes. In addition to the spatial distribution of prey, temporal patterns in prey availability and animal behavioural state were fundamental in defining risk. Our work reveals the nuanced consequences of pulsed availability on predation risk for alternative prey, whereby responses by predators to sudden resource availability, the lasting effects of diversionary prey and inherent antipredation tactics ultimately dictate risk.

A test of the frost wave hypothesis in a temperate ungulate.

Growing evidence supports the hypothesis that temperate herbivores surf the green wave of emerging plants during spring migration. Despite the importance of autumn migration, few studies have conceptualized resource tracking of temperate herbivores during this critical season. We adapted the frost wave hypothesis (FWH), which posits that animals pace their autumn migration to reduce exposure to snow but increase acquisition of forage. We tested the FWH in a population of mule deer in Wyoming, USA by tracking the autumn migrations of n = 163 mule deer that moved 15-288 km from summer to winter range. Migrating deer experienced similar amounts of snow but 1.4-2.1 times more residual forage than if they had naïve knowledge of when or how fast to migrate. Importantly, deer balanced exposure to snow and forage in a spatial manner. At the fine scale, deer avoided snow near their mountainous summer ranges and became more risk prone to snow near winter range. Aligning with their higher tolerance of snow and lingering behavior to acquire residual forage, deer increased stopover use by 1 ± 1 day (95% CI) day for every 10% of their migration completed. Our findings support the prediction that mule deer pace their autumn migration with the onset of snow and residual forage, but refine the FWH to include movement behavior en route that is spatially dynamic.

Coursing the mottled mosaic: Generalist predators track pulses in availability of neonatal ungulates.

The density and distribution of resources shape animal movement and behavior and have direct implications for population dynamics. Resource availability often is "pulsed" in space and time, and individuals should cue in on resource pulses when the energetic gain of doing so exceeds that of stable resources. Birth pulses of prey represent a profitable but ephemeral resource and should thereby result in shifting functional responses by predators. We evaluated movements and resource selection of coyotes (Canis latrans) across a gradient of reproductive stages ranging from late gestation to peak lactation of female mule deer (Odocoileus hemionus) in southwest Wyoming, USA, to test whether coyotes exhibited shifts in selection and movement behavior relative to the availability and vulnerability of neonatal mule deer. We expected coyotes to track pulses in availability of neonatal mule deer, and such behavior would be represented by shifts in resource selection and search behavior of coyotes that would be strongest during peak parturition of mule deer. Coyotes selected areas of high relative probability of use by female mule deer and did so most strongly during peak parturition. Furthermore, searching behavior of coyotes intensified during pulses of availability of deer neonates. Our findings support the notion that coyotes exploit pulses of neonatal deer, presumably as an attempt to capitalize on a vulnerable, energy-rich resource. Our work quantifies the behavioral mechanisms by which coyotes consume ungulate neonates and provides one of the first examples of a mammalian predator-prey system centered on a pulsed resource.

Migrating mule deer compensate en route for phenological mismatches.

Billions of animals migrate to track seasonal pulses in resources. Optimally timing migration is a key strategy, yet the ability of animals to compensate for phenological mismatches en route is largely unknown. Using GPS movement data collected from 72 adult female deer over a 10-year duration, we study a population of mule deer (Odocoileus hemionus) in Wyoming that lack reliable cues on their desert winter range, causing them to start migration 70 days ahead to 52 days behind the wave of spring green-up. We show that individual deer arrive at their summer range within an average 6-day window by adjusting movement speed and stopover use. Late migrants move 2.5 times faster and spend 72% less time on stopovers than early migrants, which allows them to catch the green wave. Our findings suggest that ungulates, and potentially other migratory species, possess cognitive abilities to recognize where they are in space and time relative to key resources. Such behavioral capacity may allow migratory taxa to maintain foraging benefits amid rapidly changing phenology.

Behavioural responses of a large, heat-sensitive mammal to climatic variation at multiple spatial scales.

Climate warming creates energetic challenges for endothermic species by increasing metabolic and hydric costs of thermoregulation. Although endotherms can invoke an array of behavioural and physiological strategies for maintaining homeostasis, the relative effectiveness of those strategies in a climate that is becoming both warmer and drier is not well understood. In accordance with the heat dissipation limit theory which suggests that allocation of energy to growth and reproduction by endotherms is constrained by the ability to dissipate heat, we expected that patterns of habitat use by large, heat-sensitive mammals across multiple scales are critical for behavioural thermoregulation during periods of potential heat stress and that they must invest a large portion of time to maintain heat balance. To test our predictions, we evaluated mechanisms underpinning the effectiveness of bed sites for ameliorating daytime heat loads and potential heat stress across the landscape while accounting for other factors known to affect behaviour. We integrated detailed data on microclimate and animal attributes of moose Alces alces, into a biophysical model to quantify costs of thermoregulation at fine and coarse spatial scales. During summer, moose spent an average of 67.8% of daylight hours bedded, and selected bed sites and home ranges that reduced risk of experiencing heat stress. For most of the day, shade could effectively mitigate the risk of experiencing heat stress up to 10°C, but at warmer temperatures (up to 20°C) wet soil was necessary to maintain homeostasis via conductive heat loss. Consistent selection across spatial scales for locations that reduced heat load underscores the importance of the thermal environment as a driver of behaviour in this heat-sensitive mammal. Moose in North America have long been characterized as riparian-obligate species because of their dependence on woody plant species for food. Nevertheless, the importance of dissipating endogenous heat loads conductively through wet soil suggests riparian habitats also are critical thermal refuges for moose. Such refuges may be especially important in the face of a warming climate in which both high environmental temperatures and drier conditions will likely exacerbate limits to heat dissipation, especially for large, heat-sensitive animals.

Risky business: How an herbivore navigates spatiotemporal aspects of risk from competitors and predators.

Understanding factors that influence animal behavior is central to ecology. Basic principles of animal ecology imply that individuals should seek to maximize survival and reproduction, which means carefully weighing risk against reward. Decisions become increasingly complex and constrained, however, when risk is spatiotemporally variable. We advance a growing body of work in predator-prey behavior by evaluating novel questions where a prey species is confronted with multiple predators and a potential competitor. We tested how fine-scale behavior of female mule deer (Odocoileus hemionus) during the reproductive season shifted depending upon spatial and temporal variation in risk from predators and a potential competitor. We expected female deer to avoid areas of high risk when movement activity of predators and a competitor were high. We used GPS data collected from 76 adult female mule deer, 35 adult female elk, 33 adult coyotes, and six adult mountain lions. Counter to our expectations, female deer exhibited selection for multiple risk factors, however, selection for risk was dampened by the exposure to risk within home ranges of female deer, producing a functional response in habitat selection. Furthermore, temporal variation in movement activity of predators and elk across the diel cycle did not result in a shift in movement activity by female deer. Instead, the average level of risk within their home range was the predominant factor modulating the response to risk by female deer. Our results counter prevailing hypotheses of how large herbivores navigate risky landscapes and emphasize the importance of accounting for the local environment when identifying effects of risk on animal behavior. Moreover, our findings highlight additional behavioral mechanisms used by large herbivores to mitigate multiple sources of predation and potential competitive interactions.

Biomarkers of Animal Nutrition: From Seasonal to Lifetime Indicators of Environmental Conditions.

Nutrition underpins survival and reproduction in animal populations; reliable nutritional biomarkers are therefore requisites to understanding environmental drivers of population dynamics. Biomarkers vary in scope of inference and sensitivity, making it important to know what and when to measure to properly quantify biological responses. We evaluated the repeatability of three nutritional biomarkers in a large, iteroparous mammal to evaluate the level of intrinsic and extrinsic contributions to those traits. During a long-term, individual-based study in a highly variable environment, we measured body fat, body mass, and lean mass of mule deer (Odocoileus hemionus) each autumn and spring. Lean mass was the most repeatable biomarker (0.72 autumn; 0.61 spring), followed by body mass (0.64 autumn; 0.53 spring), and then body fat (0.22 autumn; 0.01 spring). High repeatability in body and lean mass likely reflects primary structural composition, which is conserved across seasons. Low repeatability of body fat supports that it is the primary labile source of energy that is largely a product of environmental contributions of the previous season. Based on the disparate levels in repeatability among nutritional biomarkers, we contend that body and lean mass are better indicators of nutritional legacies (e.g., maternal effects), whereas body fat is a direct and sensitive reflection of recent nutritional gains and losses.

Cats and dogs: A mesopredator navigating risk and reward provisioned by an apex predator.

Successfully perceiving risk and reward is fundamental to the fitness of an animal, and can be achieved through a variety of perception tactics. For example, mesopredators may "directly" perceive risk by visually observing apex predators, or may "indirectly" perceive risk by observing habitats used by predators. Direct assessments should more accurately characterize the arrangement of risk and reward; however, indirect assessments are used more frequently in studies concerning the response of GPS-marked animals to spatiotemporally variable sources of risk and reward. We investigated the response of a mesopredator to the presence of risk and reward created by an apex predator, where risk and reward likely vary in relative perceptibility (i.e., degree of being perceptible). First, we tested whether coyotes (Canis latrans) use direct or indirect assessments to navigate the presence of mountain lions (Puma concolor; risk) and kills made by mountain lions (reward) in an area where coyotes were a common prey item for mountain lions. Second, we assessed the behavioral response of coyotes to direct encounters with mountain lions. Third, we evaluated spatiotemporal use of carrion by coyotes at kills made by mountain lions. Indirect assessments generally outperformed direct assessments when integrating analyses into a unified framework; nevertheless, our ability to detect direct perception in navigating to mountain lion kills was likely restricted by scale and sampling limitations (e.g., collar fix rates, unsampled kill sites). Rather than responding to the risk of direct encounters with mountain lions, coyotes facilitated encounters by increasing their movement rate, and engaged in risky behavior by scavenging at mountain lion kills. Coyotes appear to mitigate risk by using indirect perception to avoid mountain lions. Our predator-predator interactions and insights are nuanced and counter to the conventional predator-prey systems that have generated much of the predation risk literature.

Migration distance and maternal resource allocation determine timing of birth in a large herbivore.

Birth timing is a key life-history characteristic that influences fitness and population performance. For migratory animals, however, appropriately timing birth on one seasonal range may be constrained by events occurring during other parts of the migratory cycle. We investigated how the use of capital and income resources may facilitate flexibility in reproductive phenology of migratory mule deer in western Wyoming, USA, over a 5-yr period (2015-2019). Specifically, we examined how seasonal interactions affected three interrelated life-history characteristics: fetal development, birth mass, and birth timing. Females in good nutritional condition at the onset of winter and those that migrated short distances had more developed fetuses (measured as fetal eye diameter in March). Variation in parturition date was explained largely by fetal development; however, there were up to 16 d of plasticity in expected birth date. Plasticity in expected birth date was shaped by income resources in the form of exposure to spring green-up. Although individuals that experienced greater exposure to spring green-up were able to advance expected birth date, being born early or late with respect to fetal development had no effect on birth mass of offspring. Furthermore, we investigated the trade-offs migrating mule deer face by evaluating support for existing theory that predicts that births should be matched to local peaks in resource availability at the birth site. In contrast to this prediction, only long-distance migrants that paced migration with the flush of spring green-up, giving birth shortly after ending migration, were able to match birth with spring green-up. Shorter-distance migrants completed migration sooner and gave birth earlier, seemingly trading off more time for offspring to grow and develop over greater access to resources. Thus, movement tactic had profound downstream effects on birth timing. These findings highlight a need to reconsider classical theory on optimal birth timing, which has focused solely on conditions at the birth site.

Life-history theory provides a framework for detecting resource limitation: a test of the Nutritional Buffer Hypothesis.

For ungulates and other long-lived species, life-history theory predicts that nutritional reserves are allocated to reproduction in a state-dependent manner because survival is highly conserved. Further, as per capita food abundance and nutritional reserves decline (i.e., density dependence intensifies), reproduction and recruitment become increasingly sensitive to weather. Thus, the degree to which weather influences vital rates should be associated with proximity to nutritional carrying capacity-a notion that we refer to as the Nutritional Buffer Hypothesis. We tested the Nutritional Buffer Hypothesis using six moose (Alces alces) populations that varied in calf recruitment (33-69 calves/100 cows). We predicted that populations with high calf recruitment were nutritionally buffered against the effects of unfavorable weather, and thus were below nutritional carrying capacity. We applied a suite of tools to quantify habitat and nutritional condition of each population and found that increased browse condition, forage quality, and body fat were associated with increased pregnancy and calf recruitment, thereby providing multiple lines of evidence that declines in calf recruitment were underpinned by resource limitation. From 2001 to 2015, recruitment was more sensitive to interannual variation in weather (e.g., winter severity, drought) and plant phenology (e.g., duration of spring) for populations with reduced browse condition, forage quality, and body fat, suggesting these populations lacked the nutritional reserves necessary to buffer demographic performance against the effects of unfavorable weather. Further, average within-population calf recruitment was determined by regional climatic variation, suggesting that the pattern of reduced recruitment near the southern range boundary of moose stems from an interaction between climate and resource limitation. When coupled with information on habitat, nutrition, weather, and climate, life-history theory provides a framework to estimate nutritional limitation, proximity to nutritional carrying capacity, and impacts of climate change for ungulates.

Drivers of site fidelity in ungulates.

While the tendency to return to previously visited locations-termed 'site fidelity'-is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals' recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a 'win-stay, lose-switch' strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Linking population performance to nutritional condition in an alpine ungulate.

Bighorn sheep (Ovis canadensis) can live in extremely harsh environments and subsist on submaintenance diets for much of the year. Under these conditions, energy stored as body fat serves as an essential reserve for supplementing dietary intake to meet metabolic demands of survival and reproduction. We developed equations to predict ingesta-free body fat in bighorn sheep using ultrasonography and condition scores in vivo and carcass measurements postmortem. We then used in vivo equations to investigate the relationships between body fat, pregnancy, overwinter survival, and population growth in free-ranging bighorn sheep in California and Nevada. Among 11 subpopulations that included alpine winter residents and migrants, mean ingesta-free body fat of lactating adult females during autumn ranged between 8.8% and 15.0%; mean body fat for nonlactating females ranged from 16.4% to 20.9%. In adult females, ingesta-free body fat > 7.7% during January (early in the second trimester) corresponded with a > 90% probability of pregnancy and ingesta-free body fat > 13.5% during autumn yielded a probability of overwinter survival > 90%. Mean ingesta-free body fat of lactating females in autumn was positively associated with finite rate of population increase (λ) over the subsequent year in bighorn sheep subpopulations that wintered in alpine landscapes. Bighorn sheep with ingesta-free body fat of 26% in autumn and living in alpine environments possess energy reserves sufficient to meet resting metabolism for 83 days on fat reserves alone. We demonstrated that nutritional condition can be a pervasive mechanism underlying demography in bighorn sheep and characterizes the nutritional value of their occupied ranges. Mountain sheep are capital survivors in addition to being capital breeders, and because they inhabit landscapes with extreme seasonal forage scarcity, they also can be fat reserve obligates. Quantifying nutritional condition is essential for understanding the quality of habitats, how it underpins demography, and the proximity of a population to a nutritional threshold.

Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics.

Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called "green-wave surfing" [3-5]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Drought reshuffles plant phenology and reduces the foraging benefit of green-wave surfing for a migratory ungulate.

To increase resource gain, many herbivores pace their migration with the flush of nutritious plant green-up that progresses across the landscape (termed "green-wave surfing"). Despite concerns about the effects of climate change on migratory species and the critical role of plant phenology in mediating the ability of ungulates to surf, little is known about how drought shapes the green wave and influences the foraging benefits of migration. With a 19 year dataset on drought and plant phenology across 99 unique migratory routes of mule deer (Odocoileus hemionus) in western Wyoming, United States, we show that drought shortened the duration of spring green-up by approximately twofold (2.5 weeks) and resulted in less sequential green-up along migratory routes. We investigated the possibility that some routes were buffered from the effects of drought (i.e., routes that maintained long green-up duration irrespective of drought intensity). We found no evidence of drought-buffered routes. Instead, routes with the longest green-up in non-drought years also were the most affected by drought. Despite phenological changes along the migratory route, mule deer closely followed drought-altered green waves during migration. Migrating deer did not experience a trophic mismatch with the green wave during drought. Instead, the shorter window of green-up caused by drought reduced the opportunity to accumulate forage resources during rapid spring migrations. Our work highlights the synchronization of phenological events as an important mechanism by which climate change can negatively affect migratory species by reducing the temporal availability of key food resources. For migratory herbivores, climate change poses a new and growing threat by altering resource phenology and diminishing the foraging benefit of migration.

Chronic wasting disease undermines efforts to control the spread of brucellosis in the Greater Yellowstone Ecosystem.

Wildlife diseases pose a substantial threat to the provisioning of ecosystem services. We use a novel modeling approach to study the potential loss of these services through the imminent introduction of chronic wasting disease (CWD) to elk populations in the Greater Yellowstone Ecosystem (GYE). A specific concern is that concentrating elk at feedgrounds may exacerbate the spread of CWD, whereas eliminating feedgrounds may increase the number of elk on private ranchlands and the transmission of a second disease, brucellosis, from elk to cattle. To evaluate the consequences of management strategies given the threat of two concurrent wildlife diseases, we develop a spatiotemporal bioeconomic model. GPS data from elk and landscape attributes are used to predict migratory behavior and population densities with and without supplementary feeding. We use a 4,800 km2 area around Pinedale, Wyoming containing four existing feedgrounds as a case study. For this area, we simulate welfare estimates under a variety of management strategies. Our results indicate that continuing to feed elk could result in substantial welfare losses for the case-study region. Therefore, to maximize the present value of economic net benefits generated by the local elk population upon CWD's arrival in the region, wildlife managers may wish to consider discontinuing elk feedgrounds while simultaneously developing new methods to mitigate the financial impact to ranchers of possible brucellosis transmission to livestock. More generally, our methods can be used to weigh the costs and benefits of human-wildlife interactions in the presence of multiple disease risks.

Hunting and mountain sheep: Do current harvest practices affect horn growth?

The influence of human harvest on evolution of secondary sexual characteristics has implications for sustainable management of wildlife populations. The phenotypic consequences of selectively removing males with large horns or antlers from ungulate populations have been a topic of heightened concern in recent years. Harvest can affect size of horn-like structures in two ways: (a) shifting age structure toward younger age classes, which can reduce the mean size of horn-like structures, or (b) selecting against genes that produce large, fast-growing males. We evaluated effects of age, climatic and forage conditions, and metrics of harvest on horn size and growth of mountain sheep (Ovis canadensis ssp.) in 72 hunt areas across North America from 1981 to 2016. In 50% of hunt areas, changes in mean horn size during the study period were related to changes in age structure of harvested sheep. Environmental conditions explained directional changes in horn growth in 28% of hunt areas, 7% of which did not exhibit change before accounting for effects of the environment. After accounting for age and environment, horn size of mountain sheep was stable or increasing in the majority (~78%) of hunt areas. Age-specific horn size declined in 44% of hunt areas where harvest was regulated solely by morphological criteria, which supports the notion that harvest practices that are simultaneously selective and intensive might lead to changes in horn growth. Nevertheless, phenotypic consequences are not a foregone conclusion in the face of selective harvest; over half of the hunt areas with highly selective and intensive harvest did not exhibit age-specific declines in horn size. Our results demonstrate that while harvest regimes are an important consideration, horn growth of harvested male mountain sheep has remained largely stable, indicating that changes in horn growth patterns are an unlikely consequence of harvest across most of North America.