Growth, drought response, and climate-associated genomic structure in whitebark pine in the Sierra Nevada of California.

Whitebark pine (Pinus albicaulis Engelm.) has experienced rapid population declines and is listed as threatened under the Endangered Species Act in the United States. Whitebark pine in the Sierra Nevada of California represents the southernmost end of the species' distribution and, like other portions of its range, faces threats from an introduced pathogen, native bark beetles, and a rapidly warming climate. Beyond these chronic stressors, there is also concern about how this species will respond to acute stressors, such as drought. We present patterns of stem growth from 766 large (average diameter at breast height >25 cm), disease-free whitebark pine across the Sierra Nevada before and during a recent period of drought. We contextualize growth patterns using population genomic diversity and structure from a subset of 327 trees. Sampled whitebark pine generally had positive to neutral stem growth trends from 1970 to 2011, which was positively correlated with minimum temperature and precipitation. Indices of stem growth during drought years (2012 to 2015) relative to a predrought interval were mostly positive to neutral at our sampled sites. Individual tree growth response phenotypes appeared to be linked to genotypic variation in climate-associated loci, suggesting that some genotypes can take better advantage of local climatic conditions than others. We speculate that reduced snowpack during the 2012 to 2015 drought years may have lengthened the growing season while retaining sufficient moisture to maintain growth at most study sites. Growth responses may differ under future warming, however, particularly if drought severity increases and modifies interactions with pests and pathogens.

The energy-water limitation threshold explains divergent drought responses in tree growth, needle length, and stable isotope ratios.

Predicted increases in extreme droughts will likely cause major shifts in carbon sequestration and forest composition. Although growth declines during drought are widely documented, an increasing number of studies have reported both positive and negative responses to the same drought. These divergent growth patterns may reflect thresholds (i.e., nonlinear responses) promoted by changes in the dominant climatic constraints on tree growth. Here we tested whether stemwood growth exhibited linear or nonlinear responses to temperature and precipitation and whether stemwood growth thresholds co-occurred with multiple thresholds in source and sink processes that limit tree growth. We extracted 772 tree cores, 1398 needle length records, and 1075 stable isotope samples from 27 sites across whitebark pine's (Pinus albicaulis Engelm.) climatic niche in the Sierra Nevada. Our results indicated that a temperature threshold in stemwood growth occurred at 8.4°C (7.12-9.51°C; estimated using fall-spring maximum temperature). This threshold was significantly correlated with thresholds in foliar growth, as well as carbon (δ13 C) and nitrogen (δ15 N) stable isotope ratios, that emerged during drought. These co-occurring thresholds reflected the transition between energy- and water-limited tree growth (i.e., the E-W limitation threshold). This transition likely mediated carbon and nutrient cycling, as well as important differences in growth-defense trade-offs and drought adaptations. Furthermore, whitebark pine growing in energy-limited regions may continue to experience elevated growth in response to climate change. The positive effect of warming, however, may be offset by growth declines in water-limited regions, threatening the long-term sustainability of the recently listed whitebark pine species in the Sierra Nevada.

Nonlinear shifts in infectious rust disease due to climate change.

Range shifts of infectious plant disease are expected under climate change. As plant diseases move, emergent abiotic-biotic interactions are predicted to modify their distributions, leading to unexpected changes in disease risk. Evidence of these complex range shifts due to climate change, however, remains largely speculative. Here, we combine a long-term study of the infectious tree disease, white pine blister rust, with a six-year field assessment of drought-disease interactions in the southern Sierra Nevada. We find that climate change between 1996 and 2016 moved the climate optimum of the disease into higher elevations. The nonlinear climate change-disease relationship contributed to an estimated 5.5 (4.4-6.6) percentage points (p.p.) decline in disease prevalence in arid regions and an estimated 6.8 (5.8-7.9) p.p. increase in colder regions. Though climate change likely expanded the suitable area for blister rust by 777.9 (1.0-1392.9) km2 into previously inhospitable regions, the combination of host-pathogen and drought-disease interactions contributed to a substantial decrease (32.79%) in mean disease prevalence between surveys. Specifically, declining alternate host abundance suppressed infection probabilities at high elevations, even as climatic conditions became more suitable. Further, drought-disease interactions varied in strength and direction across an aridity gradient-likely decreasing infection risk at low elevations while simultaneously increasing infection risk at high elevations. These results highlight the critical role of aridity in modifying host-pathogen-drought interactions. Variation in aridity across topographic gradients can strongly mediate plant disease range shifts in response to climate change.

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.

Climatic stress increases forest fire severity across the western United States.

Pervasive warming can lead to chronic stress on forest trees, which may contribute to mortality resulting from fire-caused injuries. Longitudinal analyses of forest plots from across the western US show that high pre-fire climatic water deficit was related to increased post-fire tree mortality probabilities. This relationship between climate and fire was present after accounting for fire defences and injuries, and appeared to influence the effects of crown and stem injuries. Climate and fire interactions did not vary substantially across geographical regions, major genera and tree sizes. Our findings support recent physiological evidence showing that both drought and heating from fire can impair xylem conductivity. Warming trends have been linked to increasing probabilities of severe fire weather and fire spread; our results suggest that warming may also increase forest fire severity (the number of trees killed) independent of fire intensity (the amount of heat released during a fire).