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Nathan L. Stephenson

Publications and source records attributed to Nathan L. Stephenson.

At least 37 records · Page 2Linked to original sources

Compounding effects of white pine blister rust, mountain pine beetle, and fire threaten four white pine species

Invasive pathogens and bark beetles have caused precipitous declines of various tree species around the globe. Here, we characterized long‐term patterns of mountain pine beetle ( Dendroctonus ponderosae ; MPB) attacks and white pine blister rust, an infectious tree disease caused by the pathogen, Cronartium ribicola . We focused on four dominant white pine host species in Sequoia and Kings Canyon National Parks (SEKI), including sugar pine ( Pinus lambertiana ), western white pine ( P. monticola ), whitebark pine ( P. albicaulis ), and foxtail pine ( P. balfouriana ). Between 2013 and 2017, we resurveyed 152 long‐term monitoring plots that were first surveyed and established between 1995 and 1999. Overall extent (plots with at least one infected tree) of white pine blister rust (blister rust) increased from 20% to 33%. However, the infection rate across all species decreased from 5.3% to 4.2%. Blister rust dynamics varied greatly by species, as infection rate decreased from 19.1% to 6.4% in sugar pine, but increased in western white pine from 3.0% to 8.7%. For the first time, blister rust was recorded in whitebark pine, but not foxtail pine plots. MPB attacks were highest in sugar pines and decreased in the higher elevation white pine species, whitebark and foxtail pine. Both blister rust and MPB were important factors associated with elevated mortality in sugar pines. We did not, however, find a relationship between previous fires and blister rust occurrence. In addition, multiple mortality agents, including blister rust, fire, and MPB, contributed to major declines in sugar pine and western white pine; recruitment rates were much lower than mortality rates for both species. Our results highlighted that sugar pine has been declining much faster in SEKI than previously documented. If blister rust and MPB trends persist, western white pine may follow similar patterns of decline in the future. Given current spread patterns, blister rust will likely continue to increase in higher elevations, threatening subalpine white pines in the southern Sierra Nevada. More frequent long‐term monitoring efforts could inform ongoing restoration and policy focused on threats to these highly valuable and diverse white pines.

California

Mortality predispositions of conifers across western USA

Conifer mortality rates are increasing in western North America, but the physiological mechanisms underlying this trend are not well understood. We examined tree‐ring‐based radial growth along with stable carbon (C) and oxygen (O) isotope composition ( δ 13 C and δ 18 O, respectively) of dying and surviving conifers at eight old‐growth forest sites across a strong moisture gradient in the western USA to retrospectively investigate mortality predispositions. Compared with surviving trees, lower growth of dying trees was detected at least one decade before mortality at seven of the eight sites. Intrinsic water‐use efficiency increased over time in both dying and surviving trees, with a weaker increase in dying trees at five of the eight sites. C starvation was a strong correlate of conifer mortality based on a conceptual model incorporating growth, δ 13 C, and δ 18 O. However, this approach does not capture processes that occur in the final months of survival. Ultimately, C starvation may lead to increased mortality vulnerability, but hydraulic failure or biotic attack may dominate the process during the end stages of mortality in these conifers.

New Phytologist

Height-related changes in forest composition, not tree vulnerability, explain increasing mortality with height during an extreme drought

Recently, Stovall et al.1 (hereafter SSY) showed that during an extreme drought, remotely sensed mortality of tall trees was more than double that of short trees. They interpreted this to be a consequence of inherently greater hydraulic vulnerability of tall trees, and suggested that tall-tree vulnerability should thus generalize more broadly. Here we reassess their conclusions using contemporaneous, ground-based data from near their study sites. We found that 90% of trees belonged to taxonomic groups showing declining, not increasing, mortality with height, and that the overall increase in mortality with height was instead a consequence of height-related changes in forest composition, not intrinsically greater vulnerability of tall trees. Similar mechanisms likely explain mortality patterns at SSY’s sites, and, regardless, we show that SSY’s conclusions should not be accepted in the absence of robust tests of alternative mechanisms.

Nature Communications

The Fire and Tree Mortality Database, for empirical modeling of individual tree mortality after fire

Wildland fires have a multitude of ecological effects in forests, woodlands, and savannas across the globe. A major focus of past research has been on tree mortality from fire, as trees provide a vast range of biological services. We assembled a database of individual-tree records from prescribed fires and wildfires in the United States. The Fire and Tree Mortality (FTM) database includes records from 164,293 individual trees with records of fire injury (crown scorch, bole char, etc.), tree diameter, and either mortality or top-kill up to ten years post-fire. Data span 142 species and 62 genera, from 409 fires occurring from 1981-2016. Additional variables such as insect attack are included when available. The FTM database can be used to evaluate individual fire-caused mortality models for pre-fire planning and post-fire decision support, to develop improved models, and to explore general patterns of individual fire-induced tree death. The database can also be used to identify knowledge gaps that could be addressed in future research.

Scientific Data

The influence of pre-fire growth patterns on post-fire tree mortality for common conifers in western U.S. parks

Fire severity in forests is often defined in terms of post-fire tree mortality, yet the influences on tree mortality following fire are not fully understood. For trees that are not killed immediately by severe fire injury, pre-fire growth may partially predict post-fire mortality probabilities for conifers of the western U.S. Here, we consider the influence of multiple growth patterns on post-fire tree mortality. Using observations from 1 to 9 years following prescribed fires in US national parks across five western states, we show that post-fire mortality for three common conifer species is related not only to fire-caused injuries (crown scorch and bole char), but also to average growth rate and long-term (25 yr) growth patterns (counts of abrupt growth declines, and possibly growth trends). Our results suggest that pre-fire environmental and biological conditions impacting tree vigor may influence post-fire tree mortality probabilities. Fire severity, as measured by tree mortality, thus reflects tree condition as well as fire intensity. Environmental conditions (such as rising temperatures and moisture stress), independent of fire intensity, may thus cause expressed fire severity to increase in western forests.

International Journal of Wildland Fire

Size matters, but not consistently

E. Pennisi (“Forest giants are the trees most at risk,” News, 6 September, p. 962) interprets presentations of three studies as suggesting that “for trees, size is not strength, and forest giants are disproportionately vulnerable.” However, this conclusion is not well supported. The observation that lightning is a major cause of large-tree mortality on Barro Colorado Island (BCI) is best interpreted in context: when all sources of mortality are considered, small, not large BCI trees are most vulnerable to mortality (1). We have no a priori reason to assume that relative size vulnerabilities must reverse if mortality rates increase in the future. Rather than reflecting universally high drought vulnerability of large trees, the remotely-sensed observation of increasing mortality with tree height in California’s Sierra Nevada (2) likely reflects changing species dominance with height. During the drought, sizes of trees suffering greatest mortality varied widely among species, a consequence of idiosyncratic host-tree selection by different bark beetle taxa (3). Pines were the only common species with mortality that increased with size, and pines also increased in relative dominance with canopy height (4, 5). More broadly, increasing tree mortality across western USA has affected trees of all sizes (6), and a recent multi-continent compilation showed no consistent size vulnerability to drought (3). Because large trees typically are both the most highly valued and the most visible in aerial mortality surveys, the literature used to define the most damaging invasive forest pests (7) may be biased toward pests that kill large trees. Additionally, greater proportional mortality increase of large trees in the presence of invasive pests does not necessarily reflect greater absolute mortality rates relative to small trees (3, 8). Finally, Pennisi overlooked contrasting examples. For example, fire is increasing in many forests globally, and often disproportionately kills small trees (9). While large trees are certainly vulnerable to many ongoing environmental changes (10), they are not consistently the most vulnerable.

Science

Tree mortality in blue oak woodland during extreme drought in Sequoia National Park, California

Blue oak woodlands in California have been a focus of conservation concern for many years. Numerous studies have found that existing seedling and sapling numbers are inadequate to sustain current populations, and recent work has suggested that blue oak woodlands might be particularly vulnerable to a warming climate. California has recently experienced a drought of historically unprecedented severity, resulting in the mortality of tens of millions of trees, including an apparent spike in mortality in oak communities. Here we present the results of a survey of tree mortality and composition in blue oak woodlands in Sequoia National Park. We found that 18% (95% CI = 14–24,) of all standing trees and 23% (95% CI = 17–30) of standing Quercus douglasii Hook. & Arn. (blue oak) were dead, substantially higher than proportions of dead trees recorded in pre-drought datasets, which showed 4% (95% CI = 2–9) standing dead for all trees and 5% (95% CI = 4–7) dead or 8% (95% CI = 4–16) standing dead for blue oak. Furthermore, much of this mortality appeared to be recent. Based on foliage or fine twig retention, 19% (95% CI = 14–26) of blue oak and 23% (95% CI = 16–31) of Quercus wislizeni A. DC. (interior live oak) appear to have died recently. In contrast, only 5% (95% CI = 3–8) of Aesculus californica (Spach) Nutt. (California buckeye) and 5% (95% CI = 2–11) of Fraxinus dipetala Hook. & Arn. (California ash) appear to have died recently. Even after such high mortality, with blue oak basal area dropping by 26% (from 9.5 m2/ha [95% CI = 7.4–11.6] to 7.0 m2/ha [95% CI = 5.3–8.7]), blue oak remains the dominant species in these ecosystems. However, given the lack of recruitment and the apparent vulnerability to extreme drought, blue oak populations may be at risk for severe decline if such mortality events become more frequent.

California

Negative impacts of summer heat on Sierra Nevada tree seedlings

Understanding the response of forests to climate change is important for predicting changes in biodiversity and ecosystem services, including carbon storage. Seedlings represent a key demographic stage in these responses, because seedling establishment is necessary for population persistence and spread, and because the conditions allowing seedlings to survive and grow are often more restrictive than those adults can tolerate. However, detailed long‐term data on the responses of individual seedlings to climatic variation are rare. Here, we analyze a decade of seedling growth and survival data collected in 25 long‐term plots located in Sequoia and Yosemite National Parks. The forest types represented range from lower‐elevation mixed conifer, to red fir‐dominated, to high‐elevation western white pine stands. Twelve species, including two oaks and ten conifers, are found across these plots. Survival and transition to larger size classes are modeled within a Bayesian framework as a function of species, size, fire, and various climate variables. We initially expected that precipitation‐related variables would have a strong effect on seedling survival and growth in this Mediterranean‐climate region and that some species might respond positively to higher temperatures if they are adapted to warmer low‐elevation areas or to higher‐than‐average temperatures if they are high‐elevation species whose growing seasons are limited by cold. However, we found that higher July maximum temperature or temperature anomalies were associated with lower survival and growth for most species, while precipitation and snow effects were weaker and more variable. Higher precipitation had a positive effect on survival of most species, but snow effects and precipitation effects on growth could be negative. Given the estimated climate responses, under potential future scenarios of higher summer temperatures and altered precipitation, most species are predicted to have lower survival and reduced growth. This is likely to favor species that have relatively high survival and growth and less negative responses to climatic shift (such as several of the pines) and perhaps lead to a more open forest structure—or potentially transition to non‐forest vegetation types where seedling recruitment does not keep pace with adult mortality.

California

Which trees die during drought? The key role of insect host-tree selection

1. During drought, the tree subpopulations (such as size or vigor classes) that suffer disproportionate mortality can be conceptually arrayed along a continuum defined by the actions of biotic agents, particularly insects. At one extreme, stress dominates: insects are absent or simply kill the most physiologically stressed trees. At the opposite extreme, host selection dominates: outbreaking insects kill trees independently of their stress, instead selecting trees based on size or other traits. Intermediate responses are also possible. Yet for mixed-species forests, we lack a broad understanding of the relative importance of insects in determining exactly which subpopulations of trees suffer disproportionate mortality during drought, and whether these subpopulations differ among co-occurring tree species. 2. During an extreme drought, we documented the roles of native bark beetles in the mortality of five tree species in California’s Sierra Nevada. We analyzed patterns and agents of tree mortality in 12 permanent plots, and patterns of mortality in 89 temporary plots. 3. Most tree mortality was associated with bark beetles. But the growth rates (an indicator of chronic stress) and sizes of trees that suffered greatest bark-beetle-related mortality differed sharply among tree taxa, variously conforming with domination by stress (Abies concolor), domination by host selection (Pinus lambertiana and P. ponderosa), or a mix of the two (Calocedrus decurrens). Quercus kelloggii mortality remained relatively low. Thus, even during extreme drought substantial proportions of stressed trees survived because they were of sizes that mostly avoided fatal insect attack. Conversely, substantial proportions of comparatively unstressed trees died because they were of sizes that were selectively killed by outbreaking insects. 4. Synthesis. Native bark beetles were primarily responsible for determining which subpopulations of trees suffered greatest mortality during drought. However, idiosyncratic host-tree selection by the different bark beetle taxa meant that the tree subpopulations suffering greatest mortality differed strikingly among tree taxa – for example, high mortality of small trees of one species, but of large trees of another. If idiosyncratic host-tree selection by biotic mortality agents proves to be a generally common phenomenon, it could help explain weak broad-scale correlations between tree traits and tree mortality during drought.

Journal of Ecology

Pre‐fire drought and competition mediate post‐fire conifer mortality in western U.S. National Parks

Tree mortality is an important outcome of many forest fires. Extensive tree injuries from fire may lead directly to mortality, but environmental and biological stressors may also contribute to tree death. However, there is little evidence showing how the combined effects of two common stressors, drought and competition, influence post‐fire mortality. Geographically broad observations of three common western coniferous trees subjected to prescribed fire showed the likelihood of post‐fire mortality was related to intermediate‐term (10 yr) pre‐fire average radial growth, an important component of tree vigor. Path analysis showed that indices of competition and drought stress prior to fire can be described in terms of joint effects on growth, indirectly affecting post‐fire mortality. Our results suggest that the conditions that govern the relationship between growth and mortality in unburned stands may also apply to post‐fire environments. Thus, biotic and abiotic changes that affect growth negatively (e.g., drought stress) or positively (e.g., growth releases following thinning treatments) prior to fire may influence expressed fire severity, independent of fire intensity (e.g., heat flux, residence time). These relationships suggest that tree mortality may increase under stressful climatic or stand conditions even if fire behavior remains constant.

Ecological Applications

Patterns and correlates of giant sequoia foliage dieback during California’s 2012–2016 hotter drought

Hotter droughts – droughts in which unusually high temperatures exacerbate the effects of low precipitation – are expected to increase in frequency and severity in coming decades, challenging scientists and managers to identify which parts of forested landscapes may be most vulnerable. In 2014, in the middle of California’s historically unprecedented 2012–2016 hotter drought, we noticed apparently drought-induced foliage dieback in giant sequoias ( Sequoiadendron giganteum Lindl. [Buchholz]) in Sequoia and Kings Canyon national parks, California. Characteristics of the dieback were consistent with a controlled process of drought-induced senescence: younger (distal) shoots remained green while older (proximal) shoots were preferentially shed. As part of an ongoing interdisciplinary effort to understand and map sequoia vulnerability to hotter droughts, we reviewed historical records for evidence of previous foliage dieback events, surveyed dieback along trail corridors in eight sequoia groves, and analyzed tree-ring data from a high- and a low-foliage-dieback area. In sharp contrast to the greatly elevated mortality of other coniferous species found at low and middle elevations, we estimate that <1% of sequoias died during the drought. Foliage dieback was notably elevated in 2014 – the most severe single drought year in our 122-year record – but much lower in subsequent years. We found no historical records of similar foliage dieback during previous droughts. Dieback in 2014 was highly variable both within and among groves, ranging from virtually no dieback in some areas to nearly 50% in others. Dieback was highest (1) at low elevations, probably due to higher temperatures, reduced snowpack, and earlier snowmelt; (2) in areas of low adult sequoia densities, which likely reflect intrinsically more stressful sites; and (3) on steep slopes, probably reflecting reduced water availability. Average sequoia ring widths were narrower at the high-dieback than the low-dieback tree-ring site, but for reasons that remain unclear the sites did not differ in their proportional ring-width responses to past droughts. Collectively, our results suggest that giant sequoia vulnerability to hotter droughts may be spatially quite variable, and that at least some of that variability can be explained by metrics related to site water balance. Future research will focus on integrating our results with physiological and remote-sensing data, including tracking sequoias as they recover from the drought.

California

Landscape-scale variation in canopy water content of giant sequoias during drought

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.

California

Remote measurement of canopy water content in giant sequoias (Sequoiadendron giganteum) during drought

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.

California

Leaf to landscape responses of giant sequoia to hotter drought: An introduction and synthesis for the special section

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.

California

What mediates tree mortality during drought in the southern Sierra Nevada?

Severe drought has the potential to cause selective mortality within a forest, thereby inducing shifts in forest species composition. The southern Sierra Nevada foothills and mountains of California have experienced extensive forest dieback due to drought stress and insect outbreak. We used high-fidelity imaging spectroscopy (HiFIS) and light detection and ranging (LiDAR) from the Carnegie Airborne Observatory (CAO) to estimate the effect of forest dieback on species composition in response to drought stress in Sequoia National Park. Our aims were: (1) to quantify site-specific conditions that mediate tree mortality along an elevation gradient in the southern Sierra Nevada Mountains; (2) to assess where mortality events have a greater probability of occurring; and (3) to estimate which tree species have a greater likelihood of mortality along the elevation gradient. A series of statistical models were generated to classify species composition and identify tree mortality, and the influences of different environmental factors were spatially quantified and analyzed to assess where mortality events have a greater likelihood of occurring. A higher probability of mortality was observed in the lower portion of the elevation gradient, on southwest and west-facing slopes, in areas with shallow soils, on shallower slopes, and at greater distances from water. All of these factors are related to site water balance throughout the landscape. Our results also suggest that mortality is species-specific along the elevation gradient, mainly affecting Pinus ponderosa and Pinus lambertiana at lower elevations. Selective mortality within the forest may drive long-term shifts in community composition along the elevation gradient.

Sierra Nevada

Navigating translational ecology: Creating opportunities for scientist participation

Interest in translational ecology (TE) – a research approach that yields useful scientific outcomes through ongoing collaboration between scientists and stakeholders – is growing among both of these groups. Translational ecology brings together participants from different cultures and with different professional incentives. We address ways to cultivate a culture of TE, such as investing time in understanding one another's decision context and incentives, and outline common entry points to translational research, such as working through boundary organizations, building place-based research programs, and being open to opportunities as they arise. We also highlight common institutional constraints on scientists and practitioners, and ways in which collaborative research can overcome these limitations, emphasizing considerations for navigating TE within current institutional frameworks, but also pointing out ways in which institutions are evolving to facilitate translational research approaches.

Frontiers in Ecology and the Environment

Does prescribed fire promote resistance to drought in low elevation forests of the Sierra Nevada, California, USA?

Prescribed fire is a primary tool used to restore western forests following more than a century of fire exclusion, reducing fire hazard by removing dead and live fuels (small trees and shrubs). It is commonly assumed that the reduced forest density following prescribed fire also reduces competition for resources among the remaining trees, so that the remaining trees are more resistant (more likely to survive) in the face of additional stressors, such as drought. Yet this proposition remains largely untested, so that managers do not have the basic information to evaluate whether prescribed fire may help forests adapt to a future of more frequent and severe drought. During the third year of drought, in 2014, we surveyed 9950 trees in 38 burned and 18 unburned mixed conifer forest plots at low elevation (<2100 m a.s.l.) in Kings Canyon, Sequoia, and Yosemite national parks in California, USA. Fire had occurred in the burned plots from 6 yr to 28 yr before our survey. After accounting for differences in individual tree diameter, common conifer species found in the burned plots had significantly reduced probability of mortality compared to unburned plots during the drought. Stand density (stems ha -1 ) was significantly lower in burned versus unburned sites, supporting the idea that reduced competition may be responsible for the differential drought mortality response. At the time of writing, we are not sure if burned stands will maintain lower tree mortality probabilities in the face of the continued, severe drought of 2015. Future work should aim to better identify drought response mechanisms and how these may vary across other forest types and regions, particularly in other areas experiencing severe drought in the Sierra Nevada and on the Colorado Plateau.

California