Online enhancements: appendixes. Dryad data: http://dx.doi.org/10.5061/dryad.3hr2c. abstract:The forage-maturation hypothesis (FMH) states that herbivores migrate along a phenological gradient of plant development in order to maximize energy intake. Despite strong support for the FMH, the actual relationship between plant phenology and ungulate movement has remained enigmatic. We linked plant phenology (MODISnormalized difference vegetation index [NDVI] data) and space use of 167 migratory and 78 resident red deer (Cervus elaphus), using a space-time-time matrix of "springness," defined as the instantaneous rate of green-up. Consistent with the FMH, migrants experienced substantially greater access to early plant phenology than did residents. Deer were also more likely to migrate in areas where migration led to greater gains in springness. Rather than "surfing the green wave" during migration, migratory red deer moved rapidly from the winter to the summer range, thereby "jumping the green wave." However, migrants and, to a lesser degree, residents did track phenological green-up through parts of the growing season by making smaller-scale adjustments in habitat use. Despite pronounced differences in their life histories, we found only marginal differences between male and female red deer in this study. Those differences that we did detect pointed toward additional constraints on female space-use tactics, such as those posed by calving and caring for dependent offspring. We conclude that whereas in some systems migration itself is a way to surf the green wave, in others it may simply be a means to reconnect with phenological spring at the summer range. In the light of ubiquitous anthropogenic environmental change, understanding the relationship between the green wave and ungulate space use has important consequences for the management and conservation of migratory ungulates and the phenomenon of migration itself.

Partial migration is common in ungulates living in highly seasonal environments. Typically, at higher latitudes, this involves movement between high elevation summer areas used during breeding and lowland areas with less snow used during winter. Snow depth is regarded the main cause of migration to low elevation, but it is less clear why deer migrate to high elevation in spring. The forage maturation hypothesis explains the upward migration due to plant phenology. We here present also an alternative and non‐exclusive hypothesis, that deer migrate uphill in summer to escape competition due to the high density in winter areas (the competition avoidance hypothesis). We also suggest that social fences may play a role at high population density. Based on a unique study of 141 GPS‐marked red deer from seven regions covering the main distribution in Norway, we found that the proportion of migrants in the populations varied from 38% to 100%. Migration was more common in areas with a diverse topography, i.e. for areas with access to high elevation. Further, we found evidence that migration was negatively density dependent, and that fall migration was delayed at high density. We suggest that a combination of avoidance of competition in high density winter ranges, social fencing during summer in addition to the forage maturation and predation risk avoidance hypotheses, is needed to explain migration patterns of northern ungulates.

Predatory behavior of wolves (Canis lupus) was studied in 2 wolf territories in Scandinavia. We used hourly data from Global Positioning System (GPS)‐collared adult wolves in combination with Geographic Information System (GIS) for detailed analyses of movement patterns. We tested the hypothesis that wolves spend 1–2 days close to larger prey such as moose (Alces alces) and reasoned that 1–2 locations per day would be enough to find all larger prey killed by the wolves. In total, the study period comprised 287 days and yielded 6,140 hourly GPS positions, with an average of 21.4±2.4 (SD) daily positions. Depending on the radius used to define clusters, 4,045‐5,023 (65.9–81.8%) positions were included in 622–741 GPS‐clusters. We investigated all positions within clusters in the field, and 244 (22%) single positions. In total, we found 68 moose and 4 roe deer (Capreolus capreolus) and classified them as wolf‐killed within the study period. Another 10–15 moose may have been killed but not found. The GIS analyses indicated the proportion of wolf‐killed ungulates included in GPS clusters to be strongly dependent on both number of positions per day and the radius used for defining a set of spatially aggregated GPS positions as a cluster. A higher proportion (78%) of killed prey in clusters based on nighttime (2000‐0700) than those based on daytime (0800–1900) positions (41%). Simulation of aerial search during daylight hours for killed moose resulted in a serious underestimation (>60%) as compared to the number of wolf‐killed moose found during the study. The average kill rate, corrected for 14% nondetected moose, in the territories was 3.6‐4.0 days per killed moose. We concluded that the feeding behavior of wolves in Scandinavia was either different from wolves preying on moose and living at the same latitude in North America, or that estimates of wolf kill rates on moose may have been seriously underestimated in previous North American studies.