Changes in tree growth rates can affect tree mortality and forest feedbacks to the global carbon cycle. As air temperature increases, evaporative demand also increases, increasing effective drought in forest ecosystems. Using a spatially comprehensive network of Douglas fir (Pseudotsuga menziesii) chronologies from 122 locations that represent distinct climate environments in the western United States, we show that increased temperature decreases growth via vapor pressure deficit (VPD) across all latitudes. Using an ensemble of global circulation models, we project an increase in both the mean VPD associated with the lowest growth extremes and the probability of exceeding these VPD values. As temperature continues to increase in future decades, we can expect deficitrelated stress to increase and consequently Douglas fir growth to decrease throughout its US range.eclines in forest growth contribute to reduced net primary productivity and alter the global carbon cycle (1, 2). Decreased tree vigor also can predispose or increase the sensitivity of forests to disturbances and stress, such as bark beetle-induced mortality (3), related fine fuel accumulation, and short-term wildfire hazard (4), and potential mortality associated with low soil moisture (5, 6). Collectively, these changes have concomitant impacts on ecosystem processes and function (7). Changes in tree growth are often attributed to climatic variability (8, 9), wherein periods of anomalously warm and dry conditions (i.e., water stress or drought) result in decreased annual tree growth (10).Plant water stress is a combined function of water supply and demand. Increased air temperature exacerbates water stress by increasing deficits in both the soil and atmosphere. As water and atmospheric deficits increase, trees can lose water via the soilplant-atmosphere continuum, resulting in increased stress (11), or eventually close stomata, ceasing growth altogether (5). Increased precipitation can ameliorate water stress to some degree, but on seasonal and longer time scales the impact of temperature-induced increases in evaporative demand can outweigh precipitation inputs and increase water stress in forest ecosystems (12, 13).Measurements of precipitation and temperature are useful indicators of climate at any given time but, unlike "plant-relevant" variables, do not directly reflect the energy and water balance of terrestrial systems (14). Variables that express how plants "sense" climate are more useful in analyses that consider climatic limitations on plant growth and distributions (13,15). Climatic water deficit (DEF), calculated as potential evapotranspiration (PET) minus actual evapotranspiration (AET), measures climate as the interaction of water (precipitation) and energy (temperature) (14). High DEF values indicate time periods when the evaporative demand of plants is not met by available soil moisture. Vapor pressure deficit (VPD), another plant-relevant variable, is a function of relative humidity and temperature. VPD increases with temperature, and ...

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Extreme drought stress and associated bark beetle population growth contributed to an extensive tree mortality event in California, USA, resulting in more than 129 million trees dying between 2012 and 2016. Although drought is an important driver of this mortality event, past and ongoing fire suppression and the consequent densification of forests may have contributed. In some areas, land management agencies have worked to reduce stand density through mechanical treatments and prescribed fire to restore forests to less dense, more open conditions that are presumably more resilient to disturbance and changing climate. Here, we evaluate if stand structural conditions associated with treated (e.g., thinned and prescribed burned) forests in the Sierra Nevada of California conferred more resistance to the bark beetle epidemic and drought event of 2012–2016. We found that, compared to untreated units, treated units had lower stand densities, larger average tree diameters, and greater dominance of pines (Pinus), the historically dominant trees. For all tree species studied, mortality was substantially greater in climatically drier areas (i.e., lower elevations and latitudes). Both pine species studied (ponderosa pine [Pinus ponderosa] and sugar pine [Pinus lambertiana]) had greater mortality in areas where their diameters were larger, suggesting a size preference for their insect mortality agents. For ponderosa pine, the tree species experiencing greatest mortality, individual‐tree mortality probability (for a given tree diameter) was significantly lower in treated stands. Ponderosa pine mortality was also positively related to density of medium‐ to large‐sized conspecific trees, especially in areas with lower precipitation, suggesting that abundance of nearby host trees for insect mortality agents was an important determinant of pine mortality. Mortality of incense cedar (Calocedrus decurrens) and white fir (Abies concolor) was positively associated with basal area, suggesting sensitivity to competition during drought, but overall mortality was lower, likely because the most prevalent and effective mortality agents (the bark beetles Dendroctonus brevicomis and D. ponderosae) are associated specifically with pine species within our study region. Our findings suggest that forest thinning treatments are effective in reducing drought‐related tree mortality in forests, and they underscore the important interaction between water and forest density in mediating bark beetle‐caused mortality.

Historical forest conditions are often used to inform contemporary management goals because historical forests are considered to be resilient to ecological disturbances. The General Land Office (GLO) surveys of the late 19th and early 20th centuries provide regionally quasi-contiguous data sets of historical forests across much of the Western United States. Multiple methods exist for estimating tree density from point-based sampling such as the GLO surveys, including distance-based and area-based approaches. Area-based approaches have been applied in California mixed-conifer forests but their estimates have not been validated. To assess the accuracy and precision of plotless density estimators with potential for application to GLO data in this region, we imposed a GLO sampling scheme on six mapped forest stands of known densities (159-784 trees/ha) in the Sierra Nevada in California, USA, and Baja California Norte, Mexico. We compared three distance-based plotless density estimators (Cottam, Pollard, and Morisita) as well as two Voronoi area (VA) estimators, the Delincé and mean harmonic Voronoi density (MHVD), to the true densities. We simulated sampling schemes of increasing intensity to assess sampling error. The relative error (RE) of density estimates for the GLO sampling scheme ranged from 0.36 to 4.78. The least biased estimate of tree density in every stand was obtained with the Morisita estimator and the most biased was obtained with the MHVD estimator. The MHVD estimates of tree density were 1.2-3.8 times larger than the true densities and performed best in stands subject to fire exclusion for 100 yr. The Delincé approach obtained accurate estimates of density, implying that the Voronoi approach is theoretically sound but that its application in the MHVD was flawed. The misapplication was attributed to two causes: (1) the use of a crown scaling factor that does not correct for the number of trees sampled and (2) the persistent underestimate of the true VA due to a weak relationship between tree size and VA. The magnitude of differences between true densities and MHVD estimates suggest caution in using results based on the MHVD to inform management and restoration practices in the conifer forests of the American West.

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree-ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space-for-time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas-fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental | 5147 KLESSE Et aL.