Nathan L. Stephenson

CLIMATE CHANGE AND FORESTS OF THE FUTURE: MANAGING IN THE FACE OF UNCERTAINTY

We offer a conceptual framework for managing forested ecosystems under an assumption that future environments will be different from present but that we cannot be certain about the specifics of change. We encourage flexible approaches that promote reversible and incremental steps, and that favor ongoing learning and capacity to modify direction as situations change. We suggest that no single solution fits all future challenges, especially in the context of changing climates, and that the best strategy is to mix different approaches for different situations. Resources managers will be challenged to integrate adaptation strategies (actions that help ecosystems accommodate changes adaptively) and mitigation strategies (actions that enable ecosystems to reduce anthropogenic influences on global climate) into overall plans. Adaptive strategies include resistance options (forestall impacts and protect highly valued resources), resilience options (improve the capacity of ecosystems to return to desired conditions after disturbance), and response options (facilitate transition of ecosystems from current to new conditions). Mitigation strategies include options to sequester carbon and reduce overall greenhouse gas emissions. Priority-setting approaches (e.g., triage), appropriate for rapidly changing conditions and for situations where needs are greater than available capacity to respond, will become increasingly important in the future.

Widespread increase of tree mortality rates in the western United States.

Persistent changes in tree mortality rates can alter forest structure, composition, and ecosystem services such as carbon sequestration. Our analyses of longitudinal data from unmanaged old forests in the western United States showed that background (noncatastrophic) mortality rates have increased rapidly in recent decades, with doubling periods ranging from 17 to 29 years among regions. Increases were also pervasive across elevations, tree sizes, dominant genera, and past fire histories. Forest density and basal area declined slightly, which suggests that increasing mortality was not caused by endogenous increases in competition. Because mortality increased in small trees, the overall increase in mortality rates cannot be attributed solely to aging of large trees. Regional warming and consequent increases in water deficits are likely contributors to the increases in tree mortality rates.

Rate of tree carbon accumulation increases continuously with tree size

Forests are major components of the global carbon cycle, providing substantial feedback to atmospheric greenhouse gas concentrations. Our ability to understand and predict changes in the forest carbon cycle—particularly net primary productivity and carbon storage—increasingly relies on models that represent biological processes across several scales of biological organization, from tree leaves to forest stands. Yet, despite advances in our understanding of productivity at the scales of leaves and stands, no consensus exists about the nature of productivity at the scale of the individual tree, in part because we lack a broad empirical assessment of whether rates of absolute tree mass growth (and thus carbon accumulation) decrease, remain constant, or increase as trees increase in size and age. Here we present a global analysis of 403 tropical and temperate tree species, showing that for most species mass growth rate increases continuously with tree size. Thus, large, old trees do not act simply as senescent carbon reservoirs but actively fix large amounts of carbon compared to smaller trees; at the extreme, a single big tree can add the same amount of carbon to the forest within a year as is contained in an entire mid-sized tree. The apparent paradoxes of individual tree growth increasing with tree size despite declining leaf-level and stand-level productivity can be explained, respectively, by increases in a tree’s total leaf area that outpace declines in productivity per unit of leaf area and, among other factors, age-related reductions in population density. Our results resolve conflicting assumptions about the nature of tree growth, inform efforts to undertand and model forest carbon dynamics, and have additional implications for theories of resource allocation and plant senescence.

Temperate forest health in an era of emerging megadisturbance

Although disturbances such as fire and native insects can contribute to natural dynamics of forest health, exceptional droughts, directly and in combination with other disturbance factors, are pushing some temperate forests beyond thresholds of sustainability. Interactions from increasing temperatures, drought, native insects and pathogens, and uncharacteristically severe wildfire are resulting in forest mortality beyond the levels of 20th-century experience. Additional anthropogenic stressors, such as atmospheric pollution and invasive species, further weaken trees in some regions. Although continuing climate change will likely drive many areas of temperate forest toward large-scale transformations, management actions can help ease transitions and minimize losses of socially valued ecosystem services.

Tree mortality from drought, insects, and their interactions in a changing climate

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change.

REFERENCE CONDITIONS FOR GIANT SEQUOIA FOREST RESTORATION: STRUCTURE, PROCESS, AND PRECISION

National Park Service policy directs that more natural conditions be restored to giant sequoia groves, which have been altered by a century of fire exclusion. Efforts to find a reasonable and practical definition of “natural” have helped drive scientists and land managers to use past grove conditions as reference conditions for restoration. Extensive research aimed at determining reference conditions has demonstrated that past fire regimes can be characterized with greater precision than past grove structures. Difficulty and imprecision in determining past grove structure has helped fuel a debate between “structural restorationists,” who believe that forest structure should be restored mechanically before fire is reintroduced, and “process restorationists,” who believe that simple reintroduction of fire is appropriate. I evaluate old and new studies from sequoia groves to show that some of the arguments of both groups have been flawed. Importantly, it appears that restoration of fire without a preceding mechanical restoration may restore the pre-Euro-American structure of sequoia groves, at least within the bounds of our imprecise knowledge of past grove structure. However, the same may not be true for all forest types that have experienced lengthy fire exclusion. Our ability to draw robust generalizations about fires role in forest restoration will depend heavily on a thorough understanding of past and present interactions among climate, fire, and forest structure. Use of reference conditions will be central to developing this understanding.

Forest gradient response in Sierran landscapes: the physical template

Vegetation pattern on landscapes is the manifestation of physical gradients, biotic response to these gradients, and disturbances. Here we focus on the physical template as it governs the distribution of mixed-conifer forests in Californias Sierra Nevada. We extended a forest simulation model to examine montane environmental gradients, emphasizing factors affecting the water balance in these summer-dry landscapes. The model simulates the soil moisture regime in terms of the interaction of water supply and demand: supply depends on precipitation and water storage, while evapotranspirational demand varies with solar radiation and temperature. The forest cover itself can affect the water balance via canopy interception and evapotranspiration. We simulated Sierran forests as slope facets, defined as gridded stands of homogeneous topographic exposure, and verified simulated gradient response against sample quadrats distributed across Sequoia National Park. We then performed a modified sensitivity analysis of abiotic factors governing the physical gradient. Importantly, the models sensitivity to temperature, precipitation, and soil depth varies considerably over the physical template, particularly relative to elevation. The physical drivers of the water balance have characteristic spatial scales that differ by orders of magnitude. Across large spatial extents, temperature and precipitation as defined by elevation primarily govern the location of the mixed conifer zone. If the analysis is constrained to elevations within the mixed-conifer zone, local topography comes into play as it influences drainage. Soil depth varies considerably at all measured scales, and is especially dominant at fine (within-stand) scales. Physical site variables can influence soil moisture deficit either by affecting water supply or water demand; these effects have qualitatively different implications for forest response. These results have clear implications about purely inferential approaches to gradient analysis, and bear strongly on our ability to use correlative approaches in assessing the potential responses of montane forests to anthropogenic climatic change.

Forest turnover rates follow global and regional patterns of productivity

Using a global database, we found that forest turnover rates (the average of tree mortality and recruitment rates) parallel broad-scale patterns of net primary productivity. First, forest turnover was higher in tropical than in temperate forests. Second, as recently demonstrated by others, Amazonian forest turnover was higher on fertile than infertile soils. Third, within temperate latitudes, turnover was highest in angiosperm forests, intermediate in mixed forests, and lowest in gymnosperm forests. Finally, within a single forest physiognomic type, turnover declined sharply with elevation (hence with temperature). These patterns of turnover in populations of trees are broadly similar to the patterns of turnover in populations of plant organs (leaves and roots) found in other studies. Our findings suggest a link between forest mass balance and the population dynamics of trees, and have implications for understanding and predicting the effects of environmental changes on forest structure and terrestrial carbon dynamics.

Options for National parks and reserves for adapting to climate change.

Past and present climate has shaped the valued ecosystems currently protected in parks and reserves, but future climate change will redefine these conditions. Continued conservation as climate changes will require thinking differently about resource management than we have in the past; we present some logical steps and tools for doing so. Three critical tenets underpin future management plans and activities: (1) climate patterns of the past will not be the climate patterns of the future; (2) climate defines the environment and influences future trajectories of the distributions of species and their habitats; (3) specific management actions may help increase the resilience of some natural resources, but fundamental changes in species and their environment may be inevitable. Science-based management will be necessary because past experience may not serve as a guide for novel future conditions. Identifying resources and processes at risk, defining thresholds and reference conditions, and establishing monitoring and assessment programs are among the types of scientific practices needed to support a broadened portfolio of management activities. In addition to the control and hedging management strategies commonly in use today, we recommend adaptive management wherever possible. Adaptive management increases our ability to address the multiple scales at which species and processes function, and increases the speed of knowledge transfer among scientists and managers. Scenario planning provides a broad forward-thinking framework from which the most appropriate management tools can be chosen. The scope of climate change effects will require a shared vision among regional partners. Preparing for and adapting to climate change is as much a cultural and intellectual challenge as an ecological challenge.

Guiding concepts for park and wilderness stewardship in an era of global environmental change.

The major challenge to stewardship of protected areas is to decide where, when, and how to intervene in physical and biological processes, to conserve what we value in these places. To make such decisions, planners and managers must articulate more clearly the purposes of parks, what is valued, and what needs to be sustained. A key aim for conservation today is the maintenance and restoration of biodiversity, but a broader range of values are also likely to be considered important, including ecological integrity, resilience, historical fidelity (ie the ecosystem appears and functions much as it did in the past), and autonomy of nature. Until recently, the concept of “naturalness” was the guiding principle when making conservation-related decisions in park and wilderness ecosystems. However, this concept is multifaceted and often means different things to different people, including notions of historical fidelity and autonomy from human influence. Achieving the goal of nature conservation intended for such...