Drought and Predation Effects on Elk Migrations

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.


Food Webs and Migration

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. Addition- ally, 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 alternative 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.


Semi-permeable barriers and migration

1. Impermeable barriers to migration can greatly constrain the set of possible routes and ranges used by migrating animals. For ungulates, however, many forms of development are semi-permeable, and making informed management decisions about their potential impacts to the persistence of migration routes is difficult because our knowledge of how semi-permeable barriers affect migratory behaviour and function is limited.
2. Here, we propose a general framework to advance the understanding of barrier effects on ungulate migration by emphasizing the need to (i) quantify potential barriers in terms that allow behavioural thresh olds to be considered, (ii) identify and measure behavioural responses to semi-permeable barriers and (iii) consider the functional attributes of the migratory land-scape (e.g. stopover s) and how the benefits of migration might be reduced by behavioural changes.
3. We used global position system (GPS) data collected from two sub populations of mule deer Odocoileus hemionus to evaluate how different levels of gas development influenced migratory behaviour, including movement rates and stopover use at the individual level , and intensity of use and width of migration route at the population level . We then characterized the functional landscape of migration routes as either stopover habitat or movement corridors and examined how the observed behavioural changes affected the functionality of the migration route in terms of stopover use.
4. We found migratory behaviour to vary with development intensity. Our results suggest that mule deer can migrate through moderate level s of development without any notice able effects on migratory behaviour. However, in areas with more intensive development, animals often detoured from established routes, increased their rate of movement and reduced stop- over use, while the overall use and width of migration routes decreased.
5. Synthesis and applications. In contrast to impermeable barriers that impede animal movement, semi -permeable barriers allow animals to maintain connectivity between their seasonal ranges. Our results identify the mechanisms (e.g. detouring, increased movement rates, reduced stopover use) by which semi-permeable barriers affect the functionality of ungulate migration routes and emphasize that the management of semi-permeable barriers may play a key role in the conservation of migratory ungulate populations.


Crossing structures for migratory mule deer

Wildlife–vehicle collisions pose a major safety concern to motorists and can be a significant source of mortality for wildlife. Additionally, roadways can impede movements and reduce habitat connectivity. For migratory ungulates, these problems can be exacerbated when roadways bisect migration routes, as is the case in Southwest Wyoming, USA, where a 21-km section of U.S. Highway 30 overlaps with a critical winter range and migration route used by thousands of mule deer (Odocoileus hemionus). In an effort to reduce deer–vehicle collisions (DVC) and maintain migratory connectivity, the Wyoming Department of Transportation installed 7 concrete box-culvert underpasses with continuous game-proof fencing between each crossing structure. To evaluate the effectiveness of this mitigation project, we used remote cameras to quantify the number of mule deer that used the underpasses, estimate passage rates through time, and compare rates of DVCs before and after underpass construction. Through 3 years of monitoring (which encompassed autumn migration [2008, 2009, and 2010], winter use, and spring migration [2009, 2010, and 2011] for 3 migration cycles), we documented 49,146 mule deer move through the underpasses. Passage rates of deer approaching underpasses steadily increased from 54% in Year 1 to 92% in Year 3. Peak movements during the autumn migration occurred in mid-December, while peak spring movements were in mid-March and early May. Underpass and fence installation effectively reduced DVCs by 81%. Had fence gates remained closed and cattle guards clear of snow, DVCs could be eliminated altogether. Our results suggest that underpasses, combined with game-proof fencing, can improve highway safety for motorists while providing safe and effective movement corridors for large populations of migratory mule deer.


Influence of Elk Migration on Wolf Habitat Use

Identifying the ecological dynamics underlying human–wildlife conflicts is important for the management and conservation of wildlife populations. In landscapes still occupied by large carnivores, many ungulate prey species migrate seasonally, yet little empirical research has explored the relationship between carnivore distribution and ungulate migration strategy. In this study, we evaluate the influence of elk (Cervus elaphus) distribution and other landscape features on wolf ( Canis lupus ) habitat use in an area of chronic wolf– livestock conflict in the Greater Yellowstone Ecosystem, USA. Using three years of fine-scale wolf ( n ¼ 14 ) and elk ( n ¼ 81) movement data, we compared the seasonal habitat use of wolves in an area dominated by migratory elk with that of wolves in an adjacent area dominated by resident elk. Most migratory elk vacate the associated winter wolf territories each summer via a 40 – 60 km migration, whereas resident elk remain accessible to wolves year-round. We used a generalized linear model to compare the relative probability of wolf use as a function of GIS- based habitat covariates in the migratory and resident elk areas. Although wolves in both areas used elk-rich habitat all year, elk density in summer had a weaker influence on the habitat use of wolves in the migratory elk area than the resident elk area. Wolves employed a number of alternative strategies to cope with the departure of migratory elk. Wolves in the two areas also differed in their disposition toward roads. In winter, wolves in the migratory elk area used habitat close to roads, while wolves in the resident elk area avoided roads. In summer, wolves in the migratory elk area were indifferent to roads, while wolves in resident elk areas strongly avoided roads, presumably due to the location of dens and summering elk combined with different traffic levels. Study results can help wildlife managers to anticipate the movements and establishment of wolf packs as they expand into areas with migratory or resident prey populations, varying levels of human activity, and front-country rangelands with potential for conflicts with livestock.


Timing of mule deer migration

Phenological events of plants and animals are sensitive to climatic processes. Migration is a life-history event exhibited by most large herbivores living in seasonal environments, and is thought to occur in response to dynamics of forage and weather. Decisions regarding when to migrate, however, may be affected by differences in life-history characteristics of individuals. Long-term and intensive study of a population of mule deer (Odocoileus hemionus) in the Sierra Nevada, California, USA, allowed us to document patterns of migration during 11 years that encompassed a wide array of environmental conditions. We used two new techniques to properly account for interval-censored data and disentangle effects of broad-scale climate, local weather patterns, and plant phenology on seasonal patterns of migration, while incorporating effects of individual life- history characteristics. Timing of autumn migration varied substantially among individual deer, but was associated with the severity of winter weather, and in particular, snow depth and cold temperatures. Migratory responses to winter weather, however, were affected by age, nutritional condition, and summer residency of individual females. Old females and those in good nutritional condition risked encountering severe weather by delaying autumn migration, and were thus risk-prone with respect to the potential loss of foraging opportunities in deep snow compared with young females and those in poor nutritional condition. Females that summered on the west side of the crest of the Sierra Nevada delayed autumn migration relative to east-side females, which supports the influence of the local environment on timing of migration. In contrast, timing of spring migration was unrelated to individual life-history characteristics, was nearly twice as synchronous as autumn migration, differed among years, was related to the southern oscillation index, and was influenced by absolute snow depth and advancing phenology of plants. Plasticity in timing of migration in response to climatic conditions and plant phenology may be an adaptive behavioral strategy, which should reduce the detrimental effects of trophic mismatches between resources and other life-history events of large herbivores. Failure to consider effects of nutrition and other life-history traits may cloud interpretation of phenological patterns of mammals and conceal relationships associated with climate change.


Stopover ecology of a migratory ungulate

1. Birds that migrate long distances use stopover sites to optimize fuel loads and complete migration as quickly as possible. Stopover use has been predicted to facilitate a time-minimization strategy in land migrants as well, but empirical tests have been lacking, and alternative migration strategies have not been considered.
2. We used fine-scale movement data to evaluate the ecological role of stopovers in migratory mule deer Odocoileus hemionus – a land migrant whose fitness is strongly influenced by energy intake rather than migration speed.
3. Although deer could easily complete migrations (range 18–144 km) in several days, they took an average of 3 weeks and spent 95% of that time in a series of stopover sites that had higher for-age quality than movement corridors. Forage quality of stopovers increased with elevation and distance from winter range. Mule deer use of stopovers corresponded with a narrow phenological range, such that deer occupied stopovers 44 days prior to peak green-up, when forage quality was presumed to be highest. Mule deer used one stopover for every 5.3 and 6.7 km travelled during spring and autumn migrations, respectively, and used the same stopovers in consecutive years.
4. Study findings indicate that stopovers play a key role in the migration strategy of mule deer by allowing individuals to migrate in concert with plant phenology and maximize energy intake rather than speed. Our results suggest that stopover use may be more common among non-avian taxa than previously thought and, although the underlying migration strategies of temperate ungulates and birds are quite different, stopover use is important to both.
5. Exploring the role of stopovers in land migrants broadens the scope of stopover ecology and recognizes that the applied and theoretical benefits of stopover ecology need not be limited to avian taxa.


Prioritizing migration routes for conservation

As habitat loss and fragmentation increase across ungulate ranges, identifying and prioritizing migration routes for conservation has taken on new urgency. Here we present a general framework using the Brownian bridge movement model (BBMM) that: (1) provides a probabilistic estimate of the migration routes of a sampled population, (2) distinguishes between route segments that function as stopover sites vs. those used primarily as movement corridors, and (3) prioritizes routes for conservation based upon the proportion of the sampled population that uses them. We applied this approach to a migratory mule deer (Odocoileus hemionus) population in a pristine area of southwest Wyoming, USA, where 2000 gas wells and 1609 km of pipelines and roads have been proposed for development. Our analysis clearly delineated where migration routes occurred relative to proposed development and provided guidance for on-the-ground conservation efforts. Mule deer migration routes were characterized by a series of stopover sites where deer spent most of their time, connected by movement corridors through which deer moved quickly. Our findings suggest management strategies that differentiate between stopover sites and movement corridors may be warranted. Because some migration routes were used by more mule deer than others, proportional level of use may provide a reasonable metric by which routes can be prioritized for conservation. The methods we outline should be applicable to a wide range of species that inhabit regions where migration routes are threatened or poorly understood.