Assessing trends and vulnerabilities in the mutualism between whitebark pine (Pinus albicaulis) and Clark's nutcracker (Nucifraga columbiana) in national parks of the Sierra-Cascade region.

Dispersal of whitebark pine (Pinus albicaulis Engelm.), a keystone species of many high-elevation ecosystems in western North America, depends on Clark's nutcracker (Nucifraga columbiana Wilson), a seed-caching bird with an affinity for whitebark seeds. To the extent that this dependence is mutual, declines in whitebark seed production could cause declines in nutcracker abundance. Whitebark pine is in decline across much of its range due to interacting stressors, including the non-native pathogen white pine blister rust (Cronartium ribicola J. C. Fisch.). We used avian point-count data and tree surveys from four national park units to investigate whether trends in whitebark pine can explain trends in Clark's nutcracker. Spatial trends were modeled using recent data from two parks, while temporal trends were modeled using longer time-series of nutcracker and whitebark data from two additional parks. To assess the potential dependence of nutcrackers on whitebark, we linked a model of nutcracker density (accounting for detection probability) with a model of whitebark trends, using a Bayesian framework to translate uncertainty in whitebark metrics to uncertainty in nutcracker density. In Mount Rainier National Park, temporal models showed dramatic declines in nutcracker density concurrent with significant increases in whitebark crown mortality and trees infected with white pine blister rust. However, nutcrackers did not trend with whitebark metrics in North Cascades National Park Service Complex. In spatial models of data from Yosemite National Park and Sequoia-Kings Canyon National Park, nutcracker density varied not only with local cover of whitebark but also with elevation and, in Sequoia-Kings Canyon, with cover of another species of white pine. Our results add support for the hypothesis that the mutualism between whitebark pine and Clark's nutcracker is vulnerable to disruption by blister rust, and our approach integrates data across monitoring programs to explore trends in species interactions.

Landscape- and local-scale habitat influences on occurrence and detection probability of Clark's nutcrackers: Implications for conservation.

Whitebark pine (Pinus albicaulis), a keystone species and an obligate mutualist of the Clark's nutcracker (Nucifraga columbiana), is rapidly declining throughout its range. Evidence suggests this decline is leading to a downward trend in local nutcracker populations, which would in-turn decrease whitebark pine regeneration. Our objectives were to (1) evaluate temporal variation in nutcracker habitat use as a function of whitebark pine and Douglas-fir (Pseudotsuga menziesii) habitat, at local and landscape scales, (2) develop metrics for predicting when whitebark pine communities require intervention to sustain nutcracker visitation, and (3) test McKinney et al. (2009) and Barringer et al.'s (2012) models predicting nutcracker occurrence. Between 2009 and 2013, we carried out 3,135 audio-visual Clark's nutcracker surveys at 238 random points in the southern Greater Yellowstone Ecosystem. Using Bayesian occupancy models and cross-product model selection, we evaluated the association between nutcracker occurrence and habitat variables during five stages of the nutcracker annual cycle, while accounting for imperfect detection. Nutcracker occurrence was most strongly associated with the presence of cone-bearing whitebark pine trees (rather than cone density) and the area of whitebark pine on the landscape. To promote a high, >75%, probability of occurrence at a site within the study area, we recommend a management plan that achieves a landscape composed of a minimum of 12,500-25,000 ha of cone-bearing whitebark pine habitat within a 32.6 km radius. Additionally, an optimal habitat mosaic includes moderate levels of Douglas-fir habitat. Models currently used to guide whitebark pine management strategies underpredicted nutcracker occurrence in our study area, suggesting these strategies may not be appropriate in the region. We cannot predict how this mutualistic relationship will change as the population density of each species shifts. We therefore suggest conducting periodic surveys to re-evaluate the relationship as the environment changes and management strategies are implemented.

Clark's Nutcracker Breeding Season Space Use and Foraging Behavior.

Considering the entire life history of a species is fundamental to developing effective conservation strategies. Decreasing populations of five-needle white pines may be leading to the decline of Clark's nutcrackers (Nucifraga columbiana). These birds are important seed dispersers for at least ten conifer species in the western U.S., including whitebark pine (Pinus albicaulis), an obligate mutualist of Clark's nutcrackers. For effective conservation of both Clark's nutcrackers and whitebark pine, it is essential to ensure stability of Clark's nutcracker populations. My objectives were to examine Clark's nutcracker breeding season home range size, territoriality, habitat selection, and foraging behavior in the southern Greater Yellowstone Ecosystem, a region where whitebark pine is declining. I radio-tracked Clark's nutcrackers in 2011, a population-wide nonbreeding year following a low whitebark pine cone crop, and 2012, a breeding year following a high cone crop. Results suggest Douglas-fir (Pseudotsuga menziesii) communities are important habitat for Clark's nutcrackers because they selected it for home ranges. In contrast, they did not select whitebark pine habitat. However, Clark's nutcrackers did adjust their use of whitebark pine habitat between years, suggesting that, in some springs, whitebark pine habitat may be used more than previously expected. Newly extracted Douglas-fir seeds were an important food source both years. On the other hand, cached seeds made up a relatively lower proportion of the diet in 2011, suggesting cached seeds are not a reliable spring food source. Land managers focus on restoring whitebark pine habitat with the assumption that Clark's nutcrackers will be available to continue seed dispersal. In the Greater Yellowstone Ecosystem, Clark's nutcracker populations may be more likely to be retained year-round when whitebark pine restoration efforts are located adjacent to Douglas-fir habitat. By extrapolation, whitebark pine restoration efforts in other regions may consider prioritizing restoration of whitebark pine stands near alternative seed sources.

Population-Wide Failure to Breed in the Clark's Nutcracker (Nucifraga columbiana).

In highly variable environments, conditions can be so stressful in some years that entire populations forgo reproduction in favor of higher likelihood of surviving to breed in future years. In two out of five years, Clark's nutcrackers (Nucifraga Columbiana) in the Greater Yellowstone Ecosystem exhibited population-wide failure to breed. Clark's nutcrackers at the study site experienced substantial interannual differences in food availability and weather conditions, and the two nonbreeding years corresponded with low whitebark pine (Pinus albicaulis) cone crops the previous autumn (≤ an average of 8 ± 2 cones per tree versus ≥ an average of 20 ± 2 cones per tree during breeding years) and high snowpack in early spring (≥ 61.2 ± 5.5 cm versus ≤ 51.9 ± 4.4 cm during breeding years). The average adult body condition index during the breeding season was significantly lower in 2011 (-1.5 ± 1.1), a nonbreeding year, as compared to 2012 (6.2 ± 2.0), a breeding year. The environmental cues available to the birds prior to breeding, specifically availability of cached whitebark pine seeds, may have allowed them to predict that breeding conditions would be poor, leading to the decision to skip breeding. Alternatively, the Clark's nutcrackers may have had such low body energy stores that they chose not to or were unable to breed. Breeding plasticity would allow Clark's nutcrackers to exploit an unpredictable environment. However, if large-scale mortality of whitebark pines is leading to an increase in the number of nonbreeding years, there could be serious population-level and ecosystem-wide consequences.