Clinal variations in seedling traits and responses to water availability correspond to seed-source environmental gradients in a foundational dryland tree species.

BACKGROUND AND AIMS: In dryland ecosystems, conifer species are threatened by more frequent and severe droughts, which can push species beyond their physiological limits. Adequate seedling establishment will be critical for future resilience to global change. We used a common garden glasshouse experiment to determine how seedling functional trait expression and plasticity varied among seed sources in response to a gradient of water availability, focusing on a foundational dryland tree species of the western USA, Pinus monophylla. We hypothesized that the expression of growth-related seedling traits would show patterns consistent with local adaptation, given clinal variation among seed source environments. METHODS: We collected P. monophylla seeds from 23 sites distributed across rangewide gradients of aridity and seasonal moisture availability. A total of 3320 seedlings were propagated with four watering treatments representing progressively decreasing water availability. Above- and below-ground growth-related traits of first-year seedlings were measured. Trait values and trait plasticity, here representing the degree of variation among watering treatments, were modelled as a function of watering treatment and environmental conditions at the seed source locations (i.e. water availability, precipitation seasonality). KEY RESULTS: We found that, under all treatments, seedlings from more arid climates had larger above- and below-ground biomass compared to seedlings from sites experiencing lower growing-season water limitation, even after accounting for differences in seed size. Additionally, trait plasticity in response to watering treatments was greatest for seedlings from summer-wet sites that experience periodic monsoonal rain events. CONCLUSIONS: Our results show that P. monophylla seedlings respond to drought through plasticity in multiple traits, but variation in trait responses suggests that different populations are likely to respond uniquely to changes in local climate. Such trait diversity will probably influence the potential for future seedling recruitment in woodlands that are projected to experience extensive drought-related tree mortality.

Evidence of widespread topoclimatic limitation for lower treelines of the Intermountain West, United States.

Many forests in dry mountain regions are characterized by a lower elevational treeline. Understanding the controls on the position of lower treeline is important for predicting future forest distributional shifts in response to global environmental change. Lower treelines currently at their climate limit are expected to be more sensitive to changing climate, whereas lower treelines constrained by non-climatic factors are less likely to respond directly to climate change but may be sensitive to other global change agents. In this study, we used existing vegetation classifications to map lower treelines for our 1.7 million km2 study region in the U. S. Intermountain West. We modeled topoclimatic drivers of lower treeline position for each of three dominant forest types to identify topoclimatically limited treelines. We then used spatial data of edaphic properties, recent fire, and land use to identify lower treelines potentially constrained above their ecophysiological limits by non-climatic processes. We found that the lower treeline ecotone of pinyon-juniper woodlands is largely limited by topoclimate and is likely to be sensitive to increasing temperatures and associated droughts, though these effects may be heterogeneously distributed across the landscape. In contrast, dry mixed-conifer lower treelines in the northern portion of the study area rarely reached their modeled topoclimatic limit, suggesting that non-climatic processes, including fire and land use, constrain the lower treeline above its ecophysiological limits in this forest type. Our results suggest that much of the lower treeline in the Intermountain West is currently climate limited and will thus be sensitive to ongoing climate changes. Lower treelines in other arid or semi-arid mountainous regions around the globe may also be strongly sensitive to climate, though treeline response to climate change will be mediated at the local scale by soil properties, biotic interactions, and natural or anthropogenic disturbances. Our regional study of lower treeline provides a framework for identifying the drivers of lower treeline formation and allows for more robust projections of future treeline dynamics, which are needed to anticipate shifting global distributions of the forest biome.

Shrub facilitation of tree establishment varies with ontogenetic stage across environmental gradients.

Plant-plant interactions are important drivers of ecosystem structure and function, yet predicting interaction outcomes across environmental gradients remains challenging. Understanding how interactions are affected by ontogenetic shifts in plant characteristics can provide insight into the drivers of interactions and improve our ability to anticipate ecosystem responses to environmental change. We developed a conceptual framework of nurse shrub facilitation of tree establishment. We used a combination of field experiments and environmental measurements to test the framework with a shrub (Artemisia tridentata) and a tree (Pinus monophylla), two foundation species in a semiarid environment. Shrub microsites allowed trees to overcome an early population bottleneck and successfully establish in areas without tree cover. Shrubs facilitated trees at multiple ontogenetic stages, but the net outcome of the interaction shifted from strongly positive to neutral after the transition of P. monophylla from juvenile to adult foliage. Microhabitat conditions varied across a broad elevational gradient, but interaction outcomes were not strongly related to elevation. Favorable microsites provided by A. tridentata cover are crucial for P. monophylla recovery after stand-replacing disturbance. Models of vegetation response to rapid global environmental change should incorporate the critically important role of nurse shrub interactions for ameliorating population bottlenecks in tree establishment.

Fine-scale stand structure mediates drought-induced tree mortality in pinyon-juniper woodlands.

Severe drought has resulted in widespread tree die-off events in forests and woodlands globally and is forecast to become more frequent in coming decades. Tree mortality is a complex process influenced by climate, soils, characteristics of individual trees, interactions between trees, and the dynamics of pests and pathogens. The role of stand structure and stand density in mediating the resistance of trees to drought remains poorly understood, especially in semiarid woodlands, which are expected to be highly susceptible to future severe drought. We sampled permanent plots in central Nevada woodlands dominated by single-leaf pinyon pine and Utah juniper before and after a severe multi-year drought (2013-2015) to investigate the importance of climate, tree attributes, and local-neighborhood stand structure on tree mortality and canopy dieback at the level of individual trees and 0.1-ha plots. We observed widespread tree mortality of pinyon at approximately eight times the reported background mortality rate, and substantial canopy dieback in both pinyon and juniper. Both species were more prone to mortality and dieback in hotter, drier sites. Canopy dieback was associated with both long-term average climate and the severity of recent drought, with elevated mortality on sites with higher water deficits, average summer temperatures, and vapor pressure deficits. Soils also played a role in tree dieback, with greater mortality on deeper soils. While mortality was driven largely by climate at coarse scales, fine-scale stand structure interacted with climate to mediate mortality and dieback. Neighborhood statistics showed that trees were susceptible to competitive influence, and pinyon trees were especially sensitive to neighborhood density on drier sites. Mortality and dieback were associated with diverse, co-occurring insect and parasitic plant mortality agents. Canopy dieback prior to the drought was strongly associated with tree mortality during the drought, implying that current widespread defoliation caused by these agents may foreshadow future elevated woodland decline. Fine-scale influences such as stand structure and soil characteristics play a key role in the long-term dynamics of semiarid woodlands, and these factors should be considered in predictive models of forest and woodland susceptibility to drought.

Assessment of the Suitability of Tree Rings as Archives of Global and Regional Atmospheric Mercury Pollution.

This study investigated the methodology and utility of dendrochemistry in the assessment of spatial and temporal concentrations of gaseous elemental mercury. Tree cores from the Pinus species in California and Nevada, U.S.A. were collected from previously sampled areas to test the stability of tree ring concentrations over time. Cores were collected from 2 new locations to assess spatial variability among trees within and between stands located at two elevations in the same watershed. Results indicated that using 2 to 3 cores from ∼10 or more trees provided the best framework for understanding tree ring concentrations within a population of trees located in uncontaminated areas. At the least 2 sides of a tree should be cored to account for radially asymmetric variations associated with growing conditions or injury. An agreement of concentrations and trends measured in trees cored with previous research indicated that tree rings are suitable proxies for historical air mercury concentrations and that mercury concentrations have increased since the Industrial Revolution. Data collected demonstrate that tree rings record regional gradients in GEM concentrations. In addition, temporal consistency may vary within a geographic location due to differing biotic and abiotic factors influencing ring growth since trees are active samplers of atmospheric Hg.

Evidence for Nonstomatal Uptake of Hg by Aspen and Translocation of Hg from Foliage to Tree Rings in Austrian Pine.

To determine whether trees are reliable biomonitors of air mercury (Hg) pollution concentrations were measured in bark, foliage, and tree rings. Data were developed using 4-year old Pinus and Populus trees grown from common genetic stock in Oregon and subsequently transferred to four air treatments differing in gaseous oxidized mercury (GOM) chemistry and total gaseous Hg (TGM) concentrations. Soil of a subset of trees was spiked with HgBr2 in solution to test for root uptake. Results indicate no significant effect of the soil spike or GOM compounds on tree tissue Hg concentrations. TGM treatment had a significant effect on Pinus and Populus foliage, and Pinus year 5 growth ring concentrations. Populus foliar Hg concentrations were highest in the exposure where 24 h TGM concentrations were highest, indicating the importance of the nonstomatal pathway for uptake. Pinus tree ring concentrations were correlated to day time TGM concentrations suggesting Hg accumulation into tree rings is by way of the stomata and subsequent translocation by way of phloem. Populus leaves and Pinus rings can be used as biomonitors for TGM concentrations over space. However, the use of trees as temporal proxies requires further investigation due to radial translocation observed in active sapwood tree rings.

Multiscale connectivity and graph theory highlight critical areas for conservation under climate change.

Conservation planning and biodiversity management require information on landscape connectivity across a range of spatial scales from individual home ranges to large regions. Reduction in landscape connectivity due changes in land use or development is expected to act synergistically with alterations to habitat mosaic configuration arising from climate change. We illustrate a multiscale connectivity framework to aid habitat conservation prioritization in the context of changing land use and climate. Our approach, which builds upon the strengths of multiple landscape connectivity methods, including graph theory, circuit theory, and least-cost path analysis, is here applied to the conservation planning requirements of the Mohave ground squirrel. The distribution of this threatened Californian species, as for numerous other desert species, overlaps with the proposed placement of several utility-scale renewable energy developments in the American southwest. Our approach uses information derived at three spatial scales to forecast potential changes in habitat connectivity under various scenarios of energy development and climate change. By disentangling the potential effects of habitat loss and fragmentation across multiple scales, we identify priority conservation areas for both core habitat and critical corridor or stepping stone habitats. This approach is a first step toward applying graph theory to analyze habitat connectivity for species with continuously distributed habitat and should be applicable across a broad range of taxa.

Carbon dynamics in the future forest: the importance of long-term successional legacy and climate-fire interactions.

Understanding how climate change may influence forest carbon (C) budgets requires knowledge of forest growth relationships with regional climate, long-term forest succession, and past and future disturbances, such as wildfires and timber harvesting events. We used a landscape-scale model of forest succession, wildfire, and C dynamics (LANDIS-II) to evaluate the effects of a changing climate (A2 and B1 IPCC emissions; Geophysical Fluid Dynamics Laboratory General Circulation Models) on total forest C, tree species composition, and wildfire dynamics in the Lake Tahoe Basin, California, and Nevada. The independent effects of temperature and precipitation were assessed within and among climate models. Results highlight the importance of modeling forest succession and stand development processes at the landscape scale for understanding the C cycle. Due primarily to landscape legacy effects of historic logging of the Comstock Era in the late 1880s, C sequestration may continue throughout the current century, and the forest will remain a C sink (Net Ecosystem Carbon Balance > 0), regardless of climate regime. Climate change caused increases in temperatures limited simulated C sequestration potential because of augmented fire activity and reduced establishment ability of subalpine and upper montane trees. Higher temperatures influenced forest response more than reduced precipitation. As the forest reached its potential steady state, the forest could become C neutral or a C source, and climate change could accelerate this transition. The future of forest ecosystem C cycling in many forested systems worldwide may depend more on major disturbances and landscape legacies related to land use than on projected climate change alone.

The Value of Native Plants and Local Production in an Era of Global Agriculture.

For addressing potential food shortages, a fundamental tradeoff exists between investing more resources to increasing productivity of existing crops, as opposed to increasing crop diversity by incorporating more species. We explore ways to use local plants as food resources and the potential to promote food diversity and agricultural resilience. We discuss how use of local plants and the practice of local agriculture can contribute to ongoing adaptability in times of global change. Most food crops are now produced, transported, and consumed long distances from their homelands of origin. At the same time, research and practices are directed primarily at improving the productivity of a small number of existing crops that form the cornerstone of a global food economy, rather than to increasing crop diversity. The result is a loss of agro-biodiversity, leading to a food industry that is more susceptible to abiotic and biotic stressors, and more at risk of catastrophic losses. Humans cultivate only about 150 of an estimated 30,000 edible plant species worldwide, with only 30 plant species comprising the vast majority of our diets. To some extent, these practices explain the food disparity among human populations, where nearly 1 billion people suffer insufficient nutrition and 2 billion people are obese or overweight. Commercial uses of new crops and wild plants of local origin have the potential to diversify global food production and better enable local adaptation to the diverse environments humans inhabit. We discuss the advantages, obstacles, and risks of using local plants. We also describe a case study-the missed opportunity to produce pine nuts commercially in the Western United States. We discuss the potential consequences of using local pine nuts rather than importing them overseas. Finally, we provide a list of edible native plants, and synthesize the state of research concerning the potential and challenges in using them for food production. The goal of our synthesis is to support more local food production using native plants in an ecologically sustainable manner.

Spatio-temporal analysis of remotely-sensed forest mortality associated with road de-icing salts.

Forest mortality along highways has long been a concern in areas where de-icing compounds are applied during winter. This study combined the spatial advantage of high-resolution remote sensing imagery and the temporal advantage of long-term archival imagery to quantify forest mortality and to detect the subtle and chronic effects of road de-icing salts for a large mountain watershed in the Sierra Nevada Mountains, USA. IKONOS-derived mortality was used in a fine-scale spatial analysis to assess road proximity and roadside topography effects on forest mortality and to compare two potential mechanisms of de-icing salt damage, i.e. aerial deposition and soil uptake. These mechanisms were modeled using spatial proxy variables that were constructed from LiDAR topographical data. The analysis revealed a clear trend of increasing mortality with increasing potential for aerial deposition of de-icing salt onto tree crowns, mainly occurring within 10 m from roads. The effect of soil uptake of salt was weaker than that of aerial deposition but had a broader potential effect zone that extended to at least 100 m from roads. Landsat TM-derived mortality from 1989 to 2010 provided a long-term time series that indicated both immediate and lagged effects of salt application on forest mortality. Immediate effects of de-icing salt were only distinct in wet years when salt application and spray generation by passing traffic and snow plowing were likely high and other damaging factors, such as bark beetles or drought mortality, were likely weak. A strong and consistent one-year lag in the effect of salt application on incidence of mortality suggested that longer-term impacts of de-icing salt on forest health likely involved more complex pathways than simply aerial deposition. Our multi-scale remote sensing approach provided convincing evidence that de-icing salt was a significant factor for roadside forest mortality and allows for efficient future monitoring at the large-watershed scale.

Application of a rule-based model to estimate mercury exchange for three background biomes in the continental United States.

Ecosystems that have low mercury (Hg) concentrations (i.e., not enriched or impacted by geologic or anthropogenic processes) cover most of the terrestrial surface area of the earth yet their role as a net source or sink for atmospheric Hg is uncertain. Here we use empirical data to develop a rule-based model implemented within a geographic information system framework to estimate the spatial and temporal patterns of Hg flux for semiarid deserts, grasslands, and deciduous forests representing 45% of the continental United States. This exercise provides an indication of whether these ecosystems are a net source or sink for atmospheric Hg as well as a basis for recommendation of data to collect in future field sampling campaigns. Results indicated that soil alone was a small net source of atmospheric Hg and that emitted Hg could be accounted for based on Hg input by wet deposition. When foliar assimilation and wet deposition are added to the area estimate of soil Hg flux these biomes are a sink for atmospheric Hg.

Model-based assessment of aspen responses to elk herbivory in Rocky Mountain National Park, USA.

In Rocky Mountain National Park (RMNP), aspen (Populus tremuloides Michx.) has been observed to be declining on elk (Cervus elaphus nelsoni) winter range for many decades. To support elk management decisions, the SAVANNA ecosystem model was adapted to explore interactions between elk herbivory and aspen dynamics. The simulated probability of successful vegetative regeneration for senescent aspen stands declines sharply when elk densities reach levels of 3-5 elk/km2, depending on model assumptions for the seasonal duration of elk foraging activities. For aspen stands with a substantial component of younger trees, the simulated regeneration probability declines more continuously with increasing elk density, dropping below 50% from densities at 8-14 elk/km2. At the landscape scale, simulated aspen regeneration probability under a scenario of extensive seasonal use was little affected by elk population level, when this level was above 300-600 elk (25%-50% current population) over the ca. 107 km2 winter range. This was because elk distribution was highly aggregated, so that a high density of elk occupied certain areas, even at low population levels overall. At approximately current elk population levels (1000-1200 elk), only 35%-45% of senescent aspen stands are simulated as having at least a 90% probability of regeneration, nearly all of them located on the periphery of the winter range. Successful management for aspen persistence on core winter range will likely require some combination of elk population reduction, management of elk distribution, and fencing to protect aspen suckers from elk browsing.

Tree diseases and landscape processes: the challenge of landscape pathology.

Forest pathology inherently involves a landscape perspective, because tree pathogens propagate according to heterogeneous spatial patterns of flow and isolation. Landscape pathology is a field that is now emerging from the transdisciplinary cooperation of forest pathologists with landscape ecologists. Here, we review recent broad-scale assessments of tree disease risk, investigations of site and host preferences for several root rot pathogens, and regional historical analyses of pathogen outbreak in plantations. Crucial topics include fragmentation effects on pathogen spread and geophysical features that predispose forest patches to disease expression. Recent methodological developments facilitate the spatially explicit analysis of reciprocal coarse-scale relationships among hosts and pathogens. Landscape pathology studies fill a significant research gap in the context of our understanding of sustainable forest management, the introduction of exotic organisms and how climate change might affect the spread of disease.

An ecosystem approach to population management of ungulates.

Harvest objectives for wild ungulates have traditionally been based on population models that do not consider ecosystem effects of ungulate herbivory, nor interactions between native and domestic ungulate species. There is a need for ecosystem models to allow wildlife managers to evaluate potential ecosystem effects of management scenarios. The utility of the SAVANNA simulation model for estimating elk population objectives within an ecosystem context was demonstrated for North Park, Colorado, USA. Effects of different elk population levels were evaluated for range condition, elk and cattle forage, elk and cattle condition, forage and condition of mule deer and moose, plant production, and plant community composition. Analyses were based on 30-year simulation runs using variable, historical weather. Another set of analyses utilized stochastic weather patterns. For management scenarios using the historical climate pattern, increasing elk populations caused biomass reductions of palatable plant species, particularly on areas of high winter density, where mean leaf biomass of palatable shrubs declined from 26.97 g/m2 at 0 elk to 20.82 g/m2 at 4000 elk (3.73 elk/km2), a 23% decline. At population levels of 5000 elk (4.68 elk/km2) or greater, elk body condition declined sharply following a severe winter. The availability of palatable browse on critical winter range was likely the limiting factor. However, when random climate patterns were simulated for the same scenarios, the threshold level for density-dependent effects varied with climate, ranging from 2000 to 10,000 elk. We suggest that elk population levels from 4000 to 5000 animals represent a conservative population objective for the North Park elk herd. Also, increasing elk population levels appears to intensify intraspecific competition among elk, far more than interspecific competition with cattle. Resolution of elk-cattle conflicts is likely to be facilitated by managing elk distribution, rather than overall elk population levels.