Geology topics

Nathan L. Stephenson

Publications and source records attributed to Nathan L. Stephenson.

At least 19 recordsLinked to original sources

USGS2026

The state of the giant sequoias: Losses, risks, and opportunities

Background Giant sequoias, an iconic tree found only in a narrow band along the western slope of the Sierra Nevada of California, are facing unprecedented threats. Despite being adapted to fire and resistant to both drought and insect attack, the species has exhibited signs of vulnerability to these stressors in recent years, with the most substantial impacts coming from wildfire. Recently, several unprecedentedly large and severe fires have resulted in the deaths of many large trees and, in some cases, limited postfire regeneration. These impacts have led to heightened efforts by managers to implement treatments to protect those places that are still vulnerable. Here, we provide a comprehensive, range-wide assessment of the condition of the species, including an evaluation of the trends in wildfire and its effects on large giant sequoia mortality and the potential for local extirpation, treatment patterns, and vulnerability to future fire. Results From 2015 to 2024, 82% of giant sequoia grove area burned in wildfire, compared to 24% between 1910 and 2014. We conservatively estimate that 17.6% of all large giant sequoias have been killed by wildfire since 1984, with most dying in 2020 and 2021. Due to very severe wildfire, roughly 13% of the range is at some risk of local extirpation due to the loss of mature trees and limited predicted regeneration. Treatment activity has increased since 1995, with treatment type and amount varying by jurisdiction. The combination of treatment and wildfire has created a mosaic of resistance to severe fire, with 26% of grove area at high resistance, 38% at moderate resistance, 2% at low resistance, and 16% having no resistance. The remaining 18% are areas where mature giant sequoia forest was lost due to high severity fire. Conclusions Giant sequoias have experienced substantial losses due to recent wildfire, with large areas at risk of grove loss. However, most burned area had lower severity effects, creating opportunities to leverage wildfire as a treatment. When combined with areas of active management, these opportunities can promote giant sequoia conservation in a time of rapid change.

California↗

USGS2024

The relationship between maturation size and maximum tree size from tropical to boreal climates

The fundamental trade-off between current and future reproduction has long been considered to result in a tendency for species that can grow large to begin reproduction at a proportionately larger size. Due to the prolonged time required to reach maturity, estimates of tree maturation size remain unavailable and we lack a global view on the generality and the shape of this trade-off. Using seed production from five continents, we estimate tree maturation sizes for 486 tree pecies spanning tropical to boreal climates. Results show that a species’ maturation size increases with maximum size, but in a non-proportional way: the largest species begin reproduction at smaller sizes than would be expected if maturation size were simply proportional to maximum size. Furthermore, the decrease in relative maturation size is steepest in cold climates. These findings on maturation size drivers are key to accurately represent forests’ responses to disturbance and climate change.

Ecology Letters↗

USGS2024

Effects of recent wildfires on giant sequoia groves were anomalous at millennial timescales: a response to Hanson et al.

Background The giant sequoia ( Sequoiadendron giganteum [Lindley] Buchholz) of California’s Sierra Nevada recently suffered historically unprecedented wildfires that killed an estimated 13–19% of seed-bearing sequoias across their native range. Hanson et al. recently sought to characterize post-fire reproduction in two severely burned sequoia groves, but their two papers (1) inaccurately portrayed sequoia fire ecology, (2) had methodological flaws, and (3) without supporting evidence, questioned efforts to prevent large, stand-replacing wildfires and to plant sequoia seedlings in areas of low post-fire regeneration. Results Our analyses and literature review contradict many of Hanson et al.’s claims and implications. First, evidence indicates that preceding the recent wildfires, large, contiguous areas (>10 to >100 ha) of fire severe enough to kill most sequoias had been absent for at least a millennium, and probably much longer. The ancient sequoia fire regime was instead overwhelmingly dominated by surface fires in which most forest area burned at low or moderate severity interspersed with small forest gaps (hundredths of a hectare to a few hectares) created by local patches of higher-severity fire, within which most mature sequoias survived and most successful reproduction occurred. Prescribed fires have typically mimicked ancient fires and induced adequate sequoia regeneration. In contrast, in some extensive areas where recent wildfires killed most (or all) mature sequoias, regeneration has been well below historical levels, threatening a net loss of sequoia grove area. Methodologically, Hanson et al. reported sixfold greater post-fire sequoia seedling densities than others who sampled the same area; our assessments suggest their higher densities may have largely resulted from plot-placement bias. Finally, Hanson et al.’s comparisons of median seedling densities were inappropriate. Conclusions Hanson et al. questioned efforts to prevent large, high-severity wildfires in sequoia groves but did not acknowledge (1) that past fires sustained sequoia reproduction without the deaths of large fractions of mature sequoias, (2) the anomalous effects of recent wildfires, and (3) the acute conservation threat of losing large fractions of seed-bearing sequoias. Hanson et al.’s further implication, made without supporting evidence, that decisions to plant sequoia seedlings may be unwarranted ignores research showing that recent post-wildfire regeneration has often been well below historical levels.

California↗

USGS2024

Assessing giant sequoia mortality and regeneration following high-severity wildfire

Fire is a critical driver of giant sequoia ( Sequoiadendron giganteum [Lindl.] Buchholz) regeneration. However, fire suppression combined with the effects of increased temperature and severe drought has resulted in fires of an intensity and size outside of the historical norm. As a result, recent mega-fires have killed a significant portion of the world's sequoia population (13%–19%), and uncertainty surrounds whether severely affected groves will be able to recover naturally, potentially leading to a loss of grove area. To assess the likelihood of natural recovery, we collected spatially explicit data assessing mortality, crown condition, and regeneration within four giant sequoia groves that were severely impacted by the SQF- (2020) and KNP-Complex (2021) wildfires within Sequoia and Kings Canyon National Parks. In total, we surveyed 5.9 ha for seedlings and assessed the crown condition of 1104 giant sequoias. To inform management, we used a statistical methodology that robustly quantifies the uncertainty in inherently “noisy” seedling data and takes advantage of readily available remote sensing metrics that would make our findings applicable to other recently burned groves. A loss of giant sequoia grove area would be a consequence of giant sequoia tree mortality followed by a failure of natural regeneration. We found that areas that experienced very high-severity fire (above ~800 RdNBR) are at substantial risk for the loss of grove area, with tree mortality rapidly increasing and giant sequoia seedling density simultaneously decreasing with fire severity. Such high-severity areas comprised 17.8, 142.0, 14.6, 1.6 ha and ~90%, ~14%, ~53%, and ~27% of Board Camp, Redwood Mountain, Suwanee, and New Oriole Lake groves, respectively. In all sampling areas, we found that seedling densities fell far below the average density measured after prescribed fires, where seedling numbers were almost certainly adequate to maintain giant sequoia populations and postfire conditions were more in keeping with historical norms. Importantly, spatial pattern is also important in assessing the risk of grove loss, and in two groves, Suwanee and New Oriole Lake, the high-severity patches were not always contiguous, potentially making some areas more resilient to regeneration failure due to the proximity of surviving trees.

Ecosphere↗

USGS2024

Post-fire reference densities for giant sequoia seedlings in a new era of high-severity wildfires

Many forests globally are experiencing increases in large, high-severity wildfires, often with increasingly inadequate post-fire tree regeneration. To identify areas that might need post-fire planting, forest managers have a growing need for seedling reference densities – the natural seedling densities expected to be adequate to regenerate a forest – to compare with observed post-fire seedling densities. The most useful reference densities will meet five criteria: they will (1) be specific to natural post-fire reproduction rather than planted seedlings (because planted seedlings can have substantially greater survival than natural seedlings, thus underestimating adequate natural reproduction), (2) apply to the first few years following fire (when management decisions and actions are most likely), (3) be specific to each of those post-fire years (because post-fire seedling densities can change rapidly with time since fire), (4) be associated with estimates of uncertainty, and (5) include consideration of novel environmental conditions during management applications (because most reference densities will be based on data collected under more environmentally benign conditions). The world’s most massive tree species, the giant sequoia ( Sequoiadendron giganteum ) of California’s Sierra Nevada, recently experienced historically unprecedented wildfires that killed an estimated 13–19% of mature sequoias across their native range. Seedlings germinating after these fires then experienced exceptional summer heat and the two most severe summer droughts of the 121-year historical record. To help inform management responses to these events, we used seedling censuses from past fires (mostly prescribed fires) to calculate sequoia seedling reference densities meeting the five criteria. The reference densities had three striking features, which are partly attributable to giant sequoia’s status as a pioneer species. First, despite being inherently conservative, the reference densities were quite high. For example, mean first-year reference density was 172,599 seedlings ha −1 . Second, reference densities declined precipitously with time since fire: the mean fifth-year reference density was only 5% of the mean first-year density. Third, the reference densities were associated with relatively substantial uncertainty, a consequence of density variations among seedling plots; for example, the 95% credible interval for first-year reference density was 64,377 to 313,438 seedlings ha −1 . Despite this uncertainty, a case-study sequoia grove that recently burned in a high-severity wildfire had second-year post-fire seedling densities that were significantly (and dramatically) lower than the corresponding second-year reference density, suggesting inadequate post-fire reproduction. Our results highlight the value of the five criteria for reference densities – criteria that, in current practice, are rarely all met.

Forest Ecology and Management↗

USGS2023

Masting is uncommon in trees that depend on mutualist dispersers in the context of global climate and fertility gradients

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands.

Nature Plants↗

USGS2023

Snag dynamics and surface fuel loads in the Sierra Nevada: Predicting the impact of the 2012–2016 drought

Forest die-backs linked to extreme droughts are expected to increase as the climate dries and warms. An example is the 2012-2016 hotter drought in California that induced widespread tree mortality in the Sierra Nevada, California. The sudden increase in snags (i.e., standing dead trees) raised immediate concerns about their impact on wildfire hazard and longer-term questions about their impact on ecosystem structure and function. We quantified the likely progression of snag fall and fuel succession following the recent extensive mortality event in the southern Sierra Nevada mixed conifer forest. Our results used data from a long-term demography study to project trends in surface fuel loads at three study sites in Yosemite and Sequoia Kings Canyon National Parks. In the short term (2017-2021), fine woody debris and litter + duff significantly increased across all three sites (>145% and >55%, respectively); coarse woody debris increased significantly at one site (48.6%); and total fuel loads increased significantly at two of the three sites (38% and 69%). Snag longevity increased with size, with the relationship varying by species. Yellow pine was a notable outlier: size played a small role in influencing its fall rates. Overall, species-specific snag fall rates in the southern Sierra Nevada were 20% to 40% slower than previously reported. By 2040, projected median cumulative inputs of biomass from future snag fall range from 49.4 Mg ha-1 to 136.1 Mg ha-1across our three sites, which exceeds the amounts currently present (47.17-89.97 Mg ha-1) and is well above estimates of historical coarse woody debris amounts in the Sierra Nevada (17.7 Mg ha -1). These results provide a robust empirical basis to refine the snag fall algorithm in vegetation simulation models. Options to manage the impact of extreme number of snags and their large surface combustible biomass include salvage operations and prescribed burning, with both methods having operational, financial, and legal limitations that need to be considered.

Forest Ecology and Management↗

USGS2022

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

Nature Communications↗

USGS2022

Globally, tree fecundity exceeds productivity gradients

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Ecology Letters↗

USGS2022

Crowding, climate, and the case for social distancing among trees

In an emerging era of megadisturbance, bolstering forest resilience to wildfire, insects, and drought has become a central objective in many western forests. Climate has received considerable attention as a driver of these disturbances, but few studies have examined the complexities of climate–vegetation–disturbance interactions. Current strategies for creating resilient forests often rely on retrospective approaches, seeking to impart resilience by restoring historical conditions to contemporary landscapes, but historical conditions are becoming increasingly unattainable amidst modern bioclimatic conditions. What becomes an appropriate benchmark for resilience when we have novel forests, rapidly changing climate, and unprecedented disturbance regimes? We combined two longitudinal datasets—each representing some of the most comprehensive spatially explicit, annual tree mortality data in existence—in a post-hoc factorial design to examine the nonlinear relationships between fire, climate, forest spatial structure, and bark beetles. We found that while prefire drought elevated mortality risk, advantageous local neighborhoods could offset these effects. Surprisingly, mortality risk ( P m ) was higher in crowded local neighborhoods that burned in wet years ( P m = 42%) compared with sparse neighborhoods that burned during drought ( P m = 30%). Risk of beetle attack was also increased by drought, but lower conspecific crowding impeded the otherwise positive interaction between fire and beetle attack. Antecedent fire increased drought-related mortality over short timespans (<7 years) but reduced mortality over longer intervals. These results clarify interacting disturbance dynamics and provide a mechanistic underpinning for forest restoration strategies. Importantly, they demonstrate the potential for managed fire and silvicultural strategies to offset climate effects and bolster resilience to fire, beetles, and drought.

California↗

USGS2022

North American tree migration paced by climate in the West, lagging in the East

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

PNAS↗

USGS2021

Forest resistance to extended drought enhanced by prescribed fire in low elevation forests of the Sierra Nevada

Prescribed fire reduces fire hazards by removing dead and live fuels (small trees and shrubs). Reductions in forest density following prescribed fire treatments (often in concert with mechanical treatments) may also lessen competition so that residual trees might be more likely to survive when confronted with additional stressors, such as drought. The current evidence for these effects is mixed and additional study is needed. Previous work found increased tree survivorship in low elevation forests with a recent history of fire during the early years of an intense drought (2012 to 2014) in national parks in the southern Sierra Nevada. We extend these observations through additional years of intense drought and continuing elevated tree mortality through 2017 at Sequoia and Kings Canyon National Parks. Relative to unburned sites, we found that burned sites had lower stem density and had lower proportions of recently dead trees (for stems ≤47.5 cm dbh) that presumably died during the drought. Differences in recent tree mortality among burned and unburned sites held for both fir (white fir and red fir) and pine (sugar pine and ponderosa pine) species. Unlike earlier results, models of individual tree mortality probability supported an interaction between plot burn status and tree size, suggesting the effect of prescribed fire was limited to small trees. We consider differences with other recent results and discuss potential management implications including trade-offs between large tree mortality following prescribed fire and increased drought resistance.

California↗

USGS2021

Mapping the vulnerability of giant sequoias after extreme drought in California using remote sensing

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↗

USGS2021

Continent-wide tree fecundity driven by indirect climate effects

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

Nature Communications↗

USGS2021

Effects of postfire climate and seed availability on postfire conifer regeneration

Large, severe fires are becoming more frequent in many forest types across the western United States and have resulted in tree mortality across tens of thousands of hectares. Conifer regeneration in these areas is limited because seeds must travel long distances to reach the interior of large burned patches and establishment is jeopardized by increasingly hot and dry conditions. To better inform postfire management in low elevation forests of California, USA, we collected 5‐year postfire recovery data from 1,234 study plots in 19 wildfires that burned from 2004–2012 and 18 years of seed production data from 216 seed fall traps (1999–2017). We used this data in conjunction with spatially extensive estimates of climate, topography, forest composition, and burn severity to construct taxon‐specific, spatially explicit models of conifer regeneration that incorporate estimated climate conditions and seed availability during postfire recovery windows. We found that after accounting for other predictors both postfire and historical precipitation were strong predictors of regeneration, suggesting that both direct effects of postfire moisture conditions and biological inertia from historical climate may play a role in regeneration. Alternatively, postfire regeneration may simply be driven by postfire climate and apparent relationships with historical climate could be spurious. The estimated sensitivity of regeneration to postfire seed availability was strongest in firs and all conifers combined and weaker in pines. Seed production exhibited high temporal variability with seed production varying by over two orders of magnitude among years. Our models indicate that during droughts postfire conifer regeneration declines most substantially in low‐to‐moderate elevation forests. These findings enhance our mechanistic understanding of forecasted and historically documented shifts in the distribution of trees.

California↗

USGS2021

Seed production patterns of surviving Sierra Nevada conifers show minimal change following drought

Reproduction is a key component of ecological resilience in forest ecosystems, so understanding how seed production is influenced by extreme drought is key to understanding forest recovery trajectories. If trees respond to mortality-inducing drought by preferentially allocating resources for reproduction, the recovery of the stand to pre-drought conditions may be enhanced accordingly. We used a 20-year annual seed capture data set to investigate whether seed production by three tree genera commonly found in the Sierra Nevada ( Abies , Pinus , and Calocedrus ) was correlated with variation in local weather, which included an extreme drought spanning multiple years. We tested whether average seed production differed during the drought years, and whether annual seed counts could be explained by three weather variables: spring temperature, annual precipitation, and summer climatic water deficit (CWD). We fit models testing for four separate effects: (1) a priming year model (weather 1 year prior to reproductive bud initiation), (2) a bud initiation model (weather in the year of reproductive bud initiation), (3) a pollination year model (weather in the year of pollination), and (4) maturation year model (weather in the year of seed maturation). For genera with two-year reproductive cycles, the pollination and maturation models were combined. We found support for the summer CWD Abies maturation year model, which suggested higher seed outputs immediately following dry summer conditions. The spring temperature pollination year model was selected for Pinus , which suggested that seed output is higher following warm spring weather during pollination. The annual precipitation priming year model was selected for Calocedrus , which showed a negative association between seed production and wetter conditions two years prior to seed production. More parent tree basal area resulted in higher seed output for all genera, though the confidence intervals overlapped 0 for Calocedrus . Permutation tests sugested there was no systematic difference in mean seed production during the drought after accounting for live tree basal area, regardless of genus. These results highlight the variability in response across genera, and suggest that the influence of seed production on forest recovery following drought-related mortality may depend on affected species and the timing of the mortality event within the masting cycle. A greater understanding of species-level masting to drought stress is needed to more precisely predict community-level recovery following drought.

California↗

USGS2021

Why is tree drought mortality so hard to predict?

Widespread tree mortality following droughts has emerged as an environmentally and economically devastating ‘ecological surprise’. It is well established that tree physiology is important in understanding drought-driven mortality; however, the accuracy of predictions based on physiology alone has been limited. We propose that complicating factors at two levels stymie predictions of drought-driven mortality: (i) organismal-level physiological and site factors that obscure understanding of drought exposure and vulnerability and (ii) community-level ecological interactions, particularly with biotic agents whose effects on tree mortality may reverse expectations based on stress physiology. We conclude with a path forward that emphasizes the need for an integrative approach to stress physiology and biotic agent dynamics when assessing forest risk to drought-driven morality in a changing climate.

Trends in Ecology and Evolution↗

USGS2021

Preliminary estimates of sequoia mortality in the 2020 Castle Fire

Although some of California’s giant sequoia trees have stood for thousands of years and are adapted to withstand frequent low and mixed severity fires (Stephenson 1996), preliminary estimates suggest that the 2020 Castle Fire killed between 31% to 42% of large sequoias within the Castle Fire footprint, or 10% to 14% of all large sequoias across the tree’s natural range in the Sierra Nevada. This translates to an estimated loss of 7,500 to 10,600 large sequoias (those with trunk diameters of 4 feet or more). These estimates may change as new data are collected.

California↗