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.

Summary1. Well-informed management of harvested species requires understanding how changing ecological conditions affect demography and population dynamics, information that is lacking for many species. We have limited understanding of the relative influence of carnivores, harvest, weather and forage availability on elk Cervus elaphus demography, despite the ecological and economic importance of this species. We assessed adult female survival, a key vital rate for population dynamics, from 2746 radio-collared elk in 45 populations across western North America that experience wide variation in carnivore assemblage, harvest, weather and habitat conditions. 2. Proportional hazard analysis revealed that 'baseline' (i.e. not related to human factors) mortality was higher with very high winter precipitation, particularly in populations sympatric with wolves Canis lupus. Mortality may increase via nutritional stress and heightened vulnerability to predation in snowy winters. Baseline mortality was unrelated to puma Puma concolor presence, forest cover or summer forage productivity. 3. Cause-specific mortality analyses showed that wolves and all carnivore species combined had additive effects on baseline elk mortality, but only reduced survival by <2%. When human factors were included, 'total' adult mortality was solely related to harvest; the influence of native carnivores was compensatory. Annual total mortality rates were lowest in populations sympatric with both pumas and wolves because managers reduced female harvest in areas with abundant or diverse carnivores. ‡Present address: Natural Resources Canada, 506 Burnside Road W, Victoria, BC V8Z 1M5, Canada *Correspondence author. E-mail: jedediah.brodie@gmail.com †Authorship alphabetical after B. Johnson.© 2013 This article is a US Government work and is in the public domain in the USA. 2013, 50, 295-305 doi: 10.1111/1365-2664.12044 4. Mortality from native carnivores peaked in late winter and early spring, while harvest-induced mortality peaked in autumn. The strong peak in harvest-induced mortality during the autumn hunting season decreased as the number of native carnivore species increased. 5. Synthesis and applications. Elevated baseline adult female elk mortality from wolves in years with high winter precipitation could affect elk abundance as winters across the western US become drier and wolves recolonize portions of the region. In the absence of human harvest, wolves had additive, although limited, effects on mortality. However, human harvest, and its apparent use by managers to offset predation, primarily controls overall variation in adult female mortality. Altering harvest quotas is thus a strong tool for offsetting impacts of carnivore recolonization and shifting weather patterns on elk across western North America. Journal of Applied Ecology

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.

Artificial nightlight is increasingly recognized as an important environmental disturbance that influences the habitats and fitness of numerous species. However, its effects on wide‐ranging vertebrates and their interactions remain unclear. Light pollution has the potential to amplify land‐use change, and as such, answering the question of how this sensory stimulant affects behavior and habitat use of species valued for their ecological roles and economic impacts is critical for conservation and land‐use planning. Here, we combined satellite‐derived estimates of light pollution, with GPS‐data from cougars Puma concolor (n = 56), mule deer Odocoileus hemionus (n = 263) and locations of cougar‐killed deer (n = 1562 carcasses), to assess the effects of light exposure on mammal behavior and predator–prey relationships across wildland–urban gradients in the southwestern United States. Our results indicate that deer used the anthropogenic environments to access forage and were more active at night than their wildland conspecifics. Despite higher nightlight levels, cougars killed deer at the wildland–urban interface, but hunted them in the relatively darkest locations. Light had the greatest effect of all covariates on where cougars killed deer at the wildland–urban interface. Both species exhibited functional responses to light pollution at fine scales; individual cougars and deer with less light exposure increasingly avoided illuminated areas when exposed to greater radiance, whereas deer living in the wildland–urban interface selected elevated light levels. We conclude that integrating estimates of light pollution into ecological studies provides crucial insights into how the dynamic human footprint can alter animal behavior and ecosystem function across spatial scales.

Ungulates are key components in ecosystems and economically important for sport and subsistence harvest. Yet the relative importance of the effects of weather conditions, forage productivity, and carnivores on ungulates are not well understood. We examined changes in elk (Cervus canadensis) recruitment (indexed as age ratios) across 7 states and 3 ecotypes in the northwestern United States during 1989-2010, while considering the effects of predator richness, forage productivity, and precipitation. We found a broad-scale, long-term decrease in elk recruitment of 0.48 juveniles/100 adult females/year. Weather conditions (indexed as summer and winter precipitation) showed small, but measurable, influences on recruitment. Forage productivity on summer and winter ranges (indexed by normalized difference vegetation index [NDVI] metrics) had the strongest effect on elk recruitment relative to other factors. Relationships between forage productivity and recruitment varied seasonally and regionally. The productivity of winter habitat was more important in southern parts of the study area, whereas annual variation in productivity of summer habitat had more influence on recruitment in northern areas. Elk recruitment varied by up to 15 juveniles/100 adult females across the range of variation in forage productivity. Areas with more species of large carnivores had relatively low elk recruitment, presumably because of increased predation. Wolves (Canis lupus) were associated with a decrease of 5 juveniles/100 adult females, whereas grizzly bears (Ursus arctos) were associated with an additional decrease of 7 juveniles/100 adult females. Carnivore species can have a critical influence on ungulate recruitment because their influence rivals large ranges of variation in environmental conditions. A more pressing concern, however, stems from persistent broad-scale decreases in recruitment across the distribution of elk in the northwestern United States, irrespective of carnivore richness. Our results suggest that wildlife managers interested in improving recruitment of elk consider the combined effects of habitat and predators. Efforts to manage summer and winter ranges to increase forage productivity may have a positive effect on recruitment. Ó