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Increased nitrogen (N) deposition rates over the past century have affected both North American and European mountain lake ecosystems. Ecological sensitivity of mountain lakes to N deposition varies, however, because chemical and biological responses are modulated by local watershed and lake properties. We evaluated predictors of mountain lake sensitivity to atmospheric N deposition across North American and European mountain ranges and included as response variables dissolved inorganic N (DIN = N-NH4+ + N-NO3–) concentrations and phytoplankton biomass. Predictors of these responses were evaluated at three different spatial scales (hemispheric, regional, subregional) using regression tree, random forest, and generalized additive model (GAM) analysis. Analyses agreed that Northern Hemisphere mountain lake DIN was related to N deposition rates and smaller scale spatial variability (e.g., regional variability between North American and European lakes, and subregional variability between mountain ranges). Analyses suggested that DIN, N deposition, and subregional variability were important for Northern Hemisphere mountain lake phytoplankton biomass. Together, these findings highlight the need for finer-scale, subregional analyses (by mountain range) of lake sensitivity to N deposition. Subregional analyses revealed differences in predictor variables of lake sensitivity. In addition to N deposition rates, lake and watershed features such as land cover, bedrock geology, maximum lake depth (Zmax), and elevation were common modulators of lake DIN. Subregional phytoplankton biomass was consistently positively related with total phosphorus (TP) in Europe, while North American locations showed variable relationships with N or P. This study reveals scale-dependent watershed and lake characteristics modulate mountain lake ecological responses to atmospheric N deposition and provides important context to inform empirically based management strategies.

SummaryParasitic protozoa of the genusLeishmaniacycle between the phagolysosome of mammalian macrophages, where they reside as rounded intracellular amastigotes, and the midgut of female sand flies, which they colonize as elongated extracellular promastigotes. Previous studies indicated that protein kinase A (PKA) plays an important role in the initial steps of promastigote development into amastigotes. Here, we describe a novel regulatory subunit of PKA (which we have named PKAR3) that is unique toLeishmaniaand most (but not all) other Kinetoplastea. PKAR3 is localized to subpellicular microtubules (SPMT) in the cell cortex, where it recruits a specific catalytic subunit (PKAC3). Promastigotes ofPKAR3orPKAC3null mutants lose their elongated shape and are round but remain flagellated. Truncation of an N-terminal formin homology-like domain of PKAR3 results in its detachment from the SPMT, also leading to rounded promastigotes. Thus, the tethering of PKAC3 kinase activity via PKAR3 at the cell cortex is essential for maintenance of the elongated shape of promastigotes. This role of PKAR3 is reminiscent of PKARIβ and PKARIIβ binding to microtubules of mammalian neurons, which is essential for the elongation of dendrites and axons, respectively. Interestingly, PKAR3 does not bind cAMP but nucleoside analogs with a very high affinity similar to the PKAR1 isoform ofTrypanosoma. We propose that these early diverged protists have re-purposed PKA for a novel signaling pathway that spatiotemporally controls microtubule remodeling and cell shapeviaPKA activity.

Between 2012 and 2016, California suffered one of the most severe droughts on record. During this period Sequoiadendron giganteum (giant sequoias) in the Sequoia and Kings Canyon National Parks (SEKI), California, USA experienced canopy water content (CWC) loss, unprecedented foliage senescence, and, in a few cases, death. We present an assessment of the vulnerability of giant sequoia populations to droughts that is currently lacking and needed for management. We used a temporal trend of remotely sensed CWC obtained between 2015 and 2017, and recently georeferenced giant sequoia crowns to quantify the vulnerability of 7,408 individuals in 10 groves in the northern portion of SEKI. CWC is sensitive to changes in liquid water in tree canopies; therefore, it is a useful metric for quantifying the response of sequoia trees to drought. Temporal trends indicated that 9% of giant sequoias had a significant decline or consistently low CWC, suggesting these trees were likely operating at low photosynthetic capacity and potentially at high risk to drought stress. We also found that 20% of the giant sequoias had an increase or consistently high level of CWC, indicating these trees were at low risk to drought stress. These vulnerability categories were used in a random forest model with a combination of topographic, fire-related, and climate variables to generate high-resolution vulnerability risk maps. These maps show that higher risk is associated with lower elevation and higher climate water deficit. We also found that sequoias at higher elevations but located near meadows had higher vulnerability risk. These results and the vulnerability maps can identify vulnerable sequoias that may be difficult to save or locations of refugia to be protected, and thus may aid forest managers in preparation for future droughts.

California experienced severe drought from 2012 to 2016, and there were visible changes in the forest canopy throughout the State. In 2014, unprecedented foliage dieback was recorded in giant sequoia (Sequoiadendron giganteum) trees in Sequoia National Park, in the southern California Sierra Nevada mountains. Although visible changes in sequoia canopies can be recorded, biochemical and physiological responses to drought stress in giant sequoia canopies are not well understood. Ground-based measurements provide insight into the mechanisms of drought responses in trees, but are often limited to few individuals, especially in trees of tall stature such as giant sequoia. Recent studies demonstrate that remotely measured forest canopy water content (CWC) is a general indicator of canopy response to drought, but the underpinning leaf- to canopy-level causes of observed variation in CWC remain poorly understood. We combined field and airborne remote sensing measurements taken in 2015 and 2016 to assess the biophysical responses of giant sequoias to drought. In 49 study trees, CWC was related to leaf water potential, but not to the other foliar traits, suggesting that changes in CWC were made at whole-canopy rather than leaf scales. We found a non-random, spatially varying pattern in mapped CWC, with lower CWC values at lower elevation and along the outer edges of the groves. This pattern was also observed in empirical measurements of foliage dieback from the ground, and in mapped CWC across multiple sequoia groves in this region, supporting the hypothesis that drought stress is expressed in canopy-level changes in giant sequoias. The fact that we can clearly detect a relationship between CWC and foliage dieback, even without taking into account prior variability or new leaf growth, strongly suggests that remotely sensed CWC, and changes in CWC, are a useful measure of water stress in giant sequoia, and valuable for assessing and managing these iconic forests in drought.

Hotter droughts are becoming more common as climate change progresses, and they may already have caused instances of forest dieback on all forested continents. Learning from hotter droughts, including where on the landscape forests are more or less vulnerable to these events, is critical to help resource managers proactively prepare for the future. As part of our Leaf to Landscape Project, we measured the response of giant sequoia, the world’s largest tree species, to the extreme 2012–2016 hotter drought in California. The project integrated leaf-level physiology measurements, crown-level foliage dieback surveys, and remotely sensed canopy water content (CWC) to shed light on mechanisms and spatial patterns in drought response. Here we summarize initial findings, present a conceptual model of drought response, and discuss management implications; details are presented in the other four articles of the special section on Giant Sequoias and Drought. Giant sequoias exhibited both leaf- and canopy-level responses that were effective in protecting whole-tree hydraulic integrity for the vast majority of individual sequoias. Very few giant sequoias died during the drought compared to other mixed conifer tree species; however, the magnitude of sequoia drought response varied across the landscape. This variability was partially explained by local site characteristics, including variables related to site water balance. We found that low CWC is an indicator of recent foliage dieback, which occurs when stress levels are high enough that leaf-level adjustments alone are insufficient for giant sequoias to maintain hydraulic integrity. CWC or change in CWC may be useful indicators of drought stress that reveal patterns of vulnerability to future hotter droughts. Future work will measure recovery from the drought and strengthen our ability to interpret CWC maps. Our ultimate goal is to produce giant sequoia vulnerability maps to help target management actions, such as reducing other stressors, increasing resistance to hotter drought through prescribed fire or mechanical thinning, and planting sequoias in projected future suitable habitat, which may occur outside current grove distributions. We suggest that managers compare different types of vulnerability assessments and combine vulnerability maps with other sources of information to inform decisions.

Drought is expected to become an increasingly important stressor on forests globally, and understanding the physiological mechanisms driving tree drought response is essential for developing effective mitigation and conservation measures for these ecosystems. In 2014, during California’s 2012–2016 “hotter” drought in which higher temperatures exacerbated the effects of low water availability, many giant sequoia trees in the Sierra Nevada mountains exhibited foliage dieback at levels previously unreported. We hypothesized that this apparent drought-induced foliage dieback was associated with spatial patterns of site water balance and consequently tree water status and physiology. As part of an ongoing collaborative project aimed at understanding and mapping giant sequoia drought response and vulnerability at leaf to landscape scales, in 2015, 2016 and 2017 we climbed and measured leaf-level water status and physiology of mature and seedling giant sequoia trees at sites exhibiting low and high average foliage dieback in Giant Forest, Sequoia National Park. We also compared our 2015–2017 measurements with similar measurements made in giant sequoias prior to the commencement of the 2012–2016 drought. We found that during the drought, leaf water potentials of both mature and seedling sequoias were as low or lower than any previous measurements, and leaf water potentials of some seedlings were as low as the tops of mature trees. In contrast to our expectations, we found similar water potentials in both foliage dieback classes, although there was a high degree of within-site variability and in some measurement periods trees and seedlings growing in flat or meadow-side topographic positions had higher water potentials than those growing on mid- or upslope positions with presumably less favorable water supply. Leaf-level adjustments included multiple mechanisms to reduce water loss and resist the effects of desiccation at both seasonal and annual time scales, including stomatal closure, redistribution of leaf water to less mobile storage, and a shift in leaf carbon fractions to build tougher, more drought-resistant foliage. Our results suggest that giant sequoia employs a drought avoidance water regulation strategy and leaf-level adjustments are sufficient to maintain relatively isohydric water potentials above critical thresholds under most conditions. Because of the high severity of the 2012–2016 drought, however, the capacity for leaf-level compensation in some giant sequoias was overwhelmed and both leaf- and crown-level adjustments were necessary to maintain favorable water status to protect whole-tree hydraulic functioning. Additional research aimed at understanding and monitoring the consequences of future hotter droughts is necessary in order to determine the long-term persistence of giant sequoias within their current geographic distribution.

Recent drought (2012–2016) caused unprecedented foliage dieback in giant sequoias (Sequoiadendron giganteum), a species endemic to the western slope of the southern Sierra Nevada in central California. As part of an effort to understand and map sequoia response to droughts, we studied the patterns of remotely sensed canopy water content (CWC), both within and among sequoia groves in two successive years during the drought period (2015 and 2016). Our aims were: (1) to quantify giant sequoia responses to severe drought stress at a landscape scale using CWC as an indicator of crown foliage status, and (2) to estimate the effect of environmental correlates that mediate CWC change within and among giant sequoia groves. We utilized airborne high fidelity imaging spectroscopy (HiFIS) and light detection and ranging (LiDAR) data from the Carnegie Airborne Observatory to assess giant sequoia foliage status during 2015 and 2016 of the 2012–2016 droughts. A series of statistical models were generated to classify giant sequoias and to map their location in Sequoia and Kings Canyon National Parks (SEKI) and vicinity. We explored the environmental correlates and the spatial patterns of CWC change at the landscape scale. The mapped CWC was highly variable throughout the landscape during the two observation years, and proved to be most closely related to geological substrates, topography, and site-specific water balance. While there was an overall net gain in sequoia CWC between 2015 and 2016, certain locations (lower elevations, steeper slopes, areas more distant from surface water sources, and areas with greater climate water deficit) showed CWC losses. In addition, we found greater CWC loss in shorter sequoias and those growing in areas with lower sequoia stem densities. Our results suggest that CWC change indicates sequoia response to droughts across landscapes. Long-term monitoring of giant sequoia CWC will likely be useful for modeling and predicting their population-level response to future climate change.

The relative greenness and wetness of Giant Sequoia (Sequoiadendron giganteum, SEGI) groves and the surrounding Sierra Nevada, California forests were investigated using patterns in vegetation indices from Landsat imagery for the period 1985-2015. Vegetation greenness (normalized difference vegetation index) and thus forest biomass in groves increased by about 6% over that 30-year period, suggesting a 10% increase in evapotranspiration. No significant change in the surrounding non-grove forest was observed. In this period, local temperature measurements showed an increase of about 2.2 °C. The wetness of groves (normalized difference wetness index) showed no overall long-term trend, but responded to changes in annual water-year precipitation and temperature. The long-term trends of grove greenness and wetness were elevation dependent, with the lower rain-snow transition elevation zone (1700 – 2100 m) marking a change from an increasing trend at lower elevations to a decreasing trend at higher elevations. The 2011-2015 drought brought an unprecedented drop in grove wetness, over five times the 1985-2010 standard deviation; and wetness in SEGI groves dropped 50% more than in non-grove areas. Overall, the wetness and greenness of SEGI groves showed a larger response to the warming climate and drought than non-grove areas. The influence of droughts on the wetness of SEGI groves reflected effects of both the multi-decadal increase in forest biomass and the effects of warmer drought-year temperatures on the evaporative demand of current grove vegetation, plus sufficient regolith water storage of rain and snowmelt to sustain that vegetation through seasonal and multi-year dry periods.