Native forage mediates influence of irrigated agriculture on migratory behaviour of elk.

Ungulates migrate to maximize nutritional intake when forage varies seasonally. Populations of ungulates often include both migratory and non-migratory individuals, but the mechanisms driving individual differences in migratory behaviour are not well-understood. We quantified associations between hypothesized drivers of partial migration and the likelihood of migration for individual ungulates that experienced a range of environmental conditions and anthropogenic influences. We evaluated the effects of forage variation, conspecific density, and human land uses on migratory behaviour of 308 adult female elk in 16 herds across western Montana. We found irrigated agriculture on an individual's winter range reduced migratory behaviour, but individuals were more likely to migrate away from irrigated agricultural areas if better forage was available elsewhere or if they experienced high conspecific density on their winter range. When the forage available during the summer growing season varied predictably between years, elk were more likely to migrate regardless of whether they had access to irrigated agriculture. Our study shows that predictable availability of beneficial native forage can encourage migration even for ungulates with irrigated agriculture on their winter range. Perturbations that can affect the forage available to ungulates include wildfires, timber harvest, livestock grazing and changing weather patterns. If these or other disturbances negatively affect forage on summer ranges of migrants, or if they cause forage to vary unpredictably across space and time, our results suggest migratory behaviour may decline as a result.

The greenscape shapes surfing of resource waves in a large migratory herbivore.

The Green Wave Hypothesis posits that herbivore migration manifests in response to waves of spring green-up (i.e. green-wave surfing). Nonetheless, empirical support for the Green Wave Hypothesis is mixed, and a framework for understanding variation in surfing is lacking. In a population of migratory mule deer (Odocoileus hemionus), 31% surfed plant phenology in spring as well as a theoretically perfect surfer, and 98% surfed better than random. Green-wave surfing varied among individuals and was unrelated to age or energetic state. Instead, the greenscape, which we define as the order, rate and duration of green-up along migratory routes, was the primary factor influencing surfing. Our results indicate that migratory routes are more than a link between seasonal ranges, and they provide an important, but often overlooked, foraging habitat. In addition, the spatiotemporal configuration of forage resources that propagate along migratory routes shape animal movement and presumably, energy gains during migration.

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.

Foraging history of individual elephants using DNA metabarcoding.

Individual animals should adjust diets according to food availability. We used DNA metabarcoding to construct individual-level dietary timeseries for elephants from two family groups in Kenya varying in habitat use, social position and reproductive status. We detected at least 367 dietary plant taxa, with up to 137 unique plant sequences in one fecal sample. Results matched well-established trends: elephants tended to eat more grass when it rained and other plants when dry. Nested within these switches from 'grazing' to 'browsing' strategies, dietary DNA revealed seasonal shifts in food richness, composition and overlap between individuals. Elephants of both families converged on relatively cohesive diets in dry seasons but varied in their maintenance of cohesion during wet seasons. Dietary cohesion throughout the timeseries of the subdominant 'Artists' family was stronger and more consistently positive compared to the dominant 'Royals' family. The greater degree of individuality within the dominant family's timeseries could reflect more divergent nutritional requirements associated with calf dependency and/or priority access to preferred habitats. Whereas theory predicts that individuals should specialize on different foods under resource scarcity, our data suggest family bonds may promote cohesion and foster the emergence of diverse feeding cultures reflecting links between social behaviour and nutrition.

Tracking snowmelt to jump the green wave: phenological drivers of migration in a northern ungulate.

In northern climates, spring is a time of rapid environmental change: for migrating terrestrial animals, melting snow facilitates foraging and travel, and newly emergent vegetation provides a valuable nutritional resource. These changes result in selection on the timing of important life-history events such as migration and parturition occurring when high-quality resources are most abundant. We examined the timing of female caribou (Rangifer tarandus, n = 94) migration and parturition in five herds across 7 yr in Newfoundland, Canada, as a function of two measures of environmental change-snowmelt and vegetation green-up. We generated resource selection functions to test whether caribou selected for areas associated with snowmelt and green-up during migration and following calving. We found that caribou migrated approximately 1 wk prior to snowmelt, with the flush of emergent vegetation occurring during the weeks following parturition. The results indicate that caribou "jump" the green wave of emergent forage and do so by tracking the receding edge of melting snow, likely reducing movement and foraging costs related to snow cover. Our research further broadens the ecological scope of resource tracking in animals. We demonstrate that resource tracking extends beyond resources directly related to foraging to those related to movement. We also show that snowmelt provides an environmental cue that may provide a buffer against changing environmental conditions.

Complex variation in habitat selection strategies among individuals driven by extrinsic factors.

Understanding behavioral strategies employed by animals to maximize fitness in the face of environmental heterogeneity, variability, and uncertainty is a central aim of animal ecology. Flexibility in behavior may be key to how animals respond to climate and environmental change. Using a mechanistic modeling framework for simultaneously quantifying the effects of habitat preference and intrinsic movement on space use at the landscape scale, we investigate how movement and habitat selection vary among individuals and years in response to forage quality-quantity tradeoffs, environmental conditions, and variable annual climate. We evaluated the association of dynamic, biotic forage resources and static, abiotic landscape features with large grazer movement decisions in an experimental landscape, where forage resources vary in response to prescribed burning, grazing by a native herbivore, the plains bison (Bison bison bison), and a continental climate. Our goal was to determine how biotic and abiotic factors mediate bison movement decisions in a nutritionally heterogeneous grassland. We integrated spatially explicit relocations of GPS-collared bison and extensive vegetation surveys to relate movement paths to grassland attributes over a time period spanning a regionwide drought and average weather conditions. Movement decisions were affected by foliar crude content and low stature forage biomass across years with substantial interannual variation in the magnitude of selection for forage quality and quantity. These differences were associated with interannual differences in climate and growing conditions from the previous year. Our results provide experimental evidence for understanding how the forage quality-quantity tradeoff and fine-scale topography drives fine-scale movement decisions under varying environmental conditions.