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Douglas E. McWhirter

Publications and source records attributed to Douglas E. McWhirter.

11 recordsLinked to original sources

Consistent individual differences and plasticity in migration behaviour of three North American ungulates

Migratory herbivores often time spring migration to coincide with the green-up of plants. When the timing of green-up changes across years, herbivores can respond directly and be plastic to changing conditions or populations may adapt via inherent differences among individuals that may allow for an evolutionary response. We quantified plasticity and individual variation in the timing of spring migration and selection for high-quality forage as a function of the timing of spring green-up using behavioural reaction norms for three North American ungulate species. The timing of arrival to summer range (but not departure from winter range) was plastic to the timing of green-up, and both arrival and departure timing were repeatable. Our results suggest that herbivores synchronise migration with the timing of green-up by adjusting the pace of migration and may be buffered against change via individual differences. Quantifying plasticity and differences in responses represents a crucial step to elucidating the fate of species in a changing world.

Wyoming

Infection–nutrition feedbacks: Fat supports pathogen clearance but pathogens reduce fat in a wild mammal

Though far less obvious than direct effects (clinical disease or mortality), the indirect influences of pathogens are difficult to estimate but may hold fitness consequences. Here, we disentangle the directional relationships between infection and energetic reserves, evaluating the hypotheses that energetic reserves influence infection status of the host and that infection elicits costs to energetic reserves. Using repeated measures of fat reserves and infection status in individual bighorn sheep ( Ovis canadensis ) in the Greater Yellowstone Ecosystem, we documented that fat influenced ability to clear pathogens ( Mycoplasma ovipneumoniae ) and infection with respiratory pathogens was costly to fat reserves. Costs of infection approached, and in some instances exceeded, costs of rearing offspring to independence in terms of reductions to fat reserves. Fat influenced probability of clearing pathogens, pregnancy and over-winter survival; from an energetic perspective, an animal could survive for up to 23 days on the amount of fat that was lost to high levels of infection. Cost of pathogens may amplify trade-offs between reproduction and survival. In the absence of an active outbreak, the influence of resident pathogens often is overlooked. Nevertheless, the energetic burden of pathogens likely has consequences for fitness and population dynamics, especially when food resources are insufficient.

Wyoming

Forage senescence and disease influence elk pregnancy across the Greater Yellowstone Ecosystem

For various temperate ungulate species, recent research has highlighted the potential for spring vegetation phenology (“green-up”) to influence individual condition, with purported benefits to population productivity. However, few studies have been able to measure the benefit on vital rates directly, and fewer still have investigated the comparative influence of other phenological periods on ungulate vital rates. In this study, we tracked phenological changes throughout the duration of the growing season and examined how their timing affected the probability of pregnancy in an ungulate population. We did this for elk ( Cervus canadensis ) across the Greater Yellowstone Ecosystem (GYE) by sampling 1106 adult females in winter at 25 sites over a 13-year period and assessing sources of variation in pregnancy using a Bayesian hierarchical model. Pregnancy rates were generally high across the GYE (82.4%), and the primary influences on probability of pregnancy were the timing of vegetation senescence (“brown-down”) in autumn and exposure to the reproductive disease brucellosis. Earlier forage brown-down in fall negatively influenced the probability of pregnancy of elk aged 6–9 years by an estimated 17.2% within the range (ca. 32 days) of observed brown-down end dates. While summer habitat quality has been inferred to influence elk pregnancy previously, our findings specify the key influence of foraging conditions later in the seasonal cycle, immediately before the breeding season. The reproductive disease brucellosis was also an important factor, reducing the probability of pregnancy by 12.4% in elk in the 6- to 9-year age class. Because pregnancy was tested before most disease-induced abortions occur, the apparent mechanism for this effect is a prolonged reduction in fertility beyond the period of initial exposure in which fetal mortality is typically expected. Our results prompt greater scrutiny of the combined effects of late-season phenology and disease on reproductive rates and population productivity in temperate ungulates.

Idaho, Montana, Wyoming

Disease and secondary sexual traits: Effects of pneumonia on horn size of bighorn sheep

Secondary sexual traits (e.g., horns and antlers) have ecological and evolutionary importance and are of management interest for game species. Yet, how these traits respond to emerging threats like infectious disease remains underexplored. Infectious pneumonia threatens bighorn sheep ( Ovis canadensis ) populations across North America and we hypothesized it may also reduce horn growth in male sheep. We assess the effect of pneumonia on horn size in male bighorn sheep using 12 herd datasets from across the western United States that had horn growth and disease data. Disease resulted in 12–35% reduction in increment (yearly) length and 3–13% reduction in total horn length in exposed individuals. The disease effect was prolonged when pathogens continued to circulate in sheep populations. Further, disease likely delays the age at which horns reach ¾-curl and prevents achievement of full-curl. This is further evidenced with 6 of the 12 herds experiencing an increase in average age at harvest following die-off events.

Colorado, Idaho, Montana, Utah, Washington, Wyomin

Conserving transboundary wildlife migrations: Recent insights from the Greater Yellowstone Ecosystem

Animal migrations are ecologically, culturally, and economically important. Ungulate populations in many parts of Africa, Asia, Europe, and the Americas migrate long distances to access seasonally available resources, traversing vast landscapes in large numbers. Yet some migrations are declining, raising concerns among scientists and natural resource managers. We synthesize recent advances in ungulate migration ecology with relevance to management and policy. Using case studies from the Greater Yellowstone Ecosystem (GYE), we show how new tools can be applied to map ungulate migrations and assess threats across multiple seasonal habitats, serving as a conservation roadmap. To help conserve ungulate migrations, we also propose a transboundary science, policy, and management framework that could be adapted beyond the GYE and that encompasses the needs of multiple species. The key elements of this framework consist of more widespread mapping and assessment of migrations, improved federal and state coordination across jurisdictional lines, increased investment in private land conservation, and strong engagement of local stakeholders positioned to sustain conservation activities over the long term.

Idaho, Montana, Wyoming

Plasticity in elk migration timing is a response to changing environmental conditions

Migration is an effective behavioral strategy for prolonging access to seasonal resources and may be a resilient strategy for ungulates experiencing changing climatic conditions. In the Greater Yellowstone Ecosystem (GYE), elk are the primary ungulate, with approximately 20,000 individuals migrating to exploit seasonal gradients in forage while also avoiding energetically costly snow conditions. How climate-induced changes in plant phenology and snow accumulation are influencing elk migration timing is unknown. We present the most complete record of elk migration across the GYE, spanning 9 herds and 414 individuals from 2001 to 2017, to evaluate the drivers of migration timing and test for temporal shifts. The timing of elk departure from winter range involved a trade-off between current and anticipated forage conditions, while snow melt governed summer range arrival date. Timing of elk departure from summer range and arrival on winter range were both influenced by snow accumulation and exposure to hunting. At the GYE scale, spring and fall migration timing changed through time, most notably with winter range arrival dates becoming almost 50 days later since 2001. Predicted herd-level changes in migration timing largely agreed with observed GYE-wide changes—except for predicted winter range arrival dates which did not reflect the magnitude of change detected in the elk telemetry data. Snow melt, snow accumulation, and spring green-up dates all changed through time, with different herds experiencing different rates and directions of change. We conclude that elk migration is plastic, is a direct response to environmental cues, and that these environmental cues are not changing in a consistent manner across the GYE. The impacts of changing elk migration timing on predator–prey dynamics, carnivore–livestock conflict, disease ecology, and harvest management across the GYE are likely to be significant and complex.

Idaho, Montana, Wyoming

Green‐wave surfing increases fat gain in a migratory ungulate

Each spring, migratory herbivores around the world track or ‘surf’ green waves of newly emergent vegetation to distant summer or wet‐season ranges. This foraging tactic may help explain the great abundance of migratory herbivores on many seasonal landscapes. However, the underlying fitness benefits of this life‐history strategy remain poorly understood. A fundamental prediction of the green‐wave hypothesis is that migratory herbivores obtain fitness benefits from surfing waves of newly emergent vegetation more closely than their resident counterparts. Here we evaluate whether this behavior increases body‐fat levels – a critically important correlate of reproduction and survival for most ungulates – in elk Cervus elaphus of the Greater Yellowstone Ecosystem. Using satellite imagery and GPS tracking data, we found evidence that migrants (n = 23) indeed surfed the green wave, occupying sites 12.7 days closer to peak green‐up than residents (n = 16). Importantly, individual variation in surfing may help account for up to 6 kg of variation in autumn body‐fat levels. Our findings point to a pathway for anthropogenic changes to the green wave (e.g. climate change) or migrants’ ability to surf it (e.g. development) to impact migratory populations. To explore this possibility, we evaluated potential population‐level consequences of constrained surfing with a heuristic model. If green‐wave surfing deteriorates by 5–15 days from observed, our model predicts up to a 20% decrease in pregnancy rates, a 2.5% decrease in population growth, and a 30% decrease in abundance over 50 years. By linking green‐wave surfing to fitness and illustrating potential effects on population growth, our study provides new insights into the evolution of migratory behavior and the prospects for the persistence of migratory ungulate populations in a changing world.

Wyoming

Grizzly bear predation links the loss of native trout to the demography of migratory elk in Yellowstone

The loss of aquatic subsidies such as spawning salmonids is known to threaten a number of terrestrial predators, but the effects on alternative prey species are poorly understood. At the heart of the Greater Yellowstone ecosystem, an invasion of lake trout has driven a dramatic decline of native cutthroat trout that migrate up the shallow tributaries of Yellowstone Lake to spawn each spring. We explore whether this decline has amplified the effect of a generalist consumer, the grizzly bear, on populations of migratory elk that summer inside Yellowstone National Park (YNP). Recent studies of bear diets and elk populations indicate that the decline in cutthroat trout has contributed to increased predation by grizzly bears on the calves of migratory elk. Additionally, a demographic model that incorporates the increase in predation suggests that the magnitude of this diet shift has been sufficient to reduce elk calf recruitment (4–16%) and population growth (2–11%). The disruption of this aquatic–terrestrial linkage could permanently alter native species interactions in YNP. Although many recent ecological changes in YNP have been attributed to the recovery of large carnivores—particularly wolves—our work highlights a growing role of human impacts on the foraging behaviour of grizzly bears.

Yellowstone National Park

Linking anti-predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide-ranging, active predators should induce prey behavioural responses but not major, population- or community-level consequences. We evaluated the non-consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high-risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20-fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.

Montana;Wyoming;Idaho

Animal migration amid shifting patterns of phenology and predation: Lessons from a Yellowstone elk herd

Migration is a striking behavioral strategy by which many animals enhance resource acquisition while reducing predation risk. Historically, the demographic benefits of such movements made migration common, but in many taxa the phenomenon is considered globally threatened. Here we describe a long-term decline in the productivity of elk ( Cervus elaphus ) that migrate through intact wilderness areas to protected summer ranges inside Yellowstone National Park, USA. We attribute this decline to a long-term reduction in the demographic benefits that ungulates typically gain from migration. Among migratory elk, we observed a 21-year, 70% reduction in recruitment and a 4-year, 19% depression in their pregnancy rate largely caused by infrequent reproduction of females that were young or lactating. In contrast, among resident elk, we have recently observed increasing recruitment and a high rate of pregnancy. Landscape-level changes in habitat quality and predation appear to be responsible for the declining productivity of Yellowstone migrants. From 1989 to 2009, migratory elk experienced an increasing rate and shorter duration of green-up coincident with warmer spring–summer temperatures and reduced spring precipitation, also consistent with observations of an unusually severe drought in the region. Migrants are also now exposed to four times as many grizzly bears ( Ursus arctos ) and wolves ( Canis lupus ) as resident elk. Both of these restored predators consume migratory elk calves at high rates in the Yellowstone wilderness but are maintained at low densities via lethal management and human disturbance in the year-round habitats of resident elk. Our findings suggest that large-carnivore recovery and drought, operating simultaneously along an elevation gradient, have disproportionately influenced the demography of migratory elk. Many migratory animals travel large geographic distances between their seasonal ranges. Changes in land use and climate that disparately influence such seasonal ranges may alter the ecological basis of migratory behavior, representing an important challenge for, and a powerful lens into, the ecology and conservation of migratory taxa.

Idaho, Montana, Wyoming

Rejoinder: Challenge and opportunity in the study of ungulate migration amid environmental change

Increasingly, animals that migrate long distances to exploit seasonal habitats must traverse political boundaries capable of altering the very ecological gradients that promote migratory behavior. This transboundary aspect of migration presents many new challenges and opportunities for research and conservation (e.g., Bolger et al. 2008, Taillon et al. 2012). Work to date has often focused on physical barriers to movement (roads, fences,and housing and energy development) that can threaten migratory populations to varying degrees (Holdo et al. 2011, Sawyer et al. 2013). However, even in the absence of conspicuous barriers, political and jurisdictional boundaries can bring dramatic differences in land use and conservation policy. What happens to migratory populations when these boundaries alter the resources and refuges that they seek on their seasonal journeys?

Ecology