Mark E. Harmon

Ecology of Coarse Woody Debris in Temperate Ecosystems

Publisher Summary This chapter reviews the rates at which Coarse Woody Debris (CWD) is added and removed from ecosystems, the biomass found in streams and forests, and many functions that CWD serves. CWD is an important component of temperate stream and forest ecosystems and is added to the ecosystem by numerous mechanisms, including wind, fire, insect attack, pathogens, competition, and geomorphic processes. Many factors control the rate at which CWD decomposes, including temperature, moisture, the internal gas composition of CWD, substrate quality, the size of the CWD, and the types of organisms involved. The mass of CWD in an ecosystem ideally represents the balance between addition and loss. In reality, slow decomposition rates and erratic variations in input of CWD cause the CWD mass to deviate markedly from steady-state projections. Many differences correspond to forest type, with deciduous-dominated systems having generally lower biomass than conifer-dominated systems. Stream size also influences CWD mass in lotic ecosystems, while successional stage dramatically influences CWD mass in boat aquatic and terrestrial settings. This chapter reviews many of these functions and concludes that CWD is an important functional component of stream and forest ecosystems. Better scientific understanding of these functions and the natural factors influencing CWD dynamics should lead to more enlightened management practices.

Disturbances and structural development of natural forest ecosystems with silvicultural implications, using Douglas-fir forests as an example

Forest managers need a comprehensive scientific understanding of natural stand development processes when designing silvicultural systems that integrate ecological and economic objectives, including a better appreciation of the nature of disturbance regimes and the biological legacies, such as live trees, snags, and logs, that they leave behind. Most conceptual forest development models do not incorporate current knowledge of the: (1) complexity of structures (including spatial patterns) and developmental processes; (2) duration of development in long-lived forests; (3) complex spatial patterns of stands that develop in later stages of seres; and particularly (4) the role of disturbances in creating structural legacies that become key elements of the post-disturbance stands. We elaborate on existing models for stand structural development using natural stand development of the Douglas-fir—western hemlock sere in the Pacific Northwest as our primary example; most of the principles are broadly applicable while some processes (e.g. role of epicormic branches) are related to specific species. We discuss the use of principles from disturbance ecology and natural stand development to create silvicultural approaches that are more aligned with natural processes. Such approaches provide for a greater abundance of standing dead and down wood and large old trees, perhaps reducing short-term commercial productivity but ultimately enhancing wildlife habitat, biodiversity, and ecosystem function, including soil protection and nutrient retention. # 2002 Elsevier Science B.V. All rights reserved.

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.

Tree Death as an Ecological Process

the patterns and causes of tree death typically are complex, and we are only beginning to appreciate the complexities. Understanding and predicting tree mortality is critical in both applied and basic ecology. Practically speaking, information on mortality is essential in calculating forest stand yields and allocating efforts in tending and protecting forests. A thorough knowledge of tree death is also necessary to interpret correctly the dying back of forests. Yet, despite its long history, forest husbandry lacks a comprehensive understanding of tree mortality. In basic ecology, tree death is relevant to a broad array of topics. Ecolo-

Effects on carbon storage of conversion of old-growth forests to young forests.

Simulations of carbon storage suggest that conversion of old-growth forests to young fast-growing forests will not decrease atmospheric carbon dioxide (CO2) in general, as has been suggested recently. During simulated timber harvest, on-site carbon storage is reduced considerably and does not approach old-growth storage capacity for at least 200 years. Even when sequestration of carbon in wooden buildings is included in the models, timber harvest results in a net flux of CO2 to the atmosphere. To offset this effect, the production of lumber and other long-term wood products, as well as the life-span of buildings, would have to increase markedly. Mass balance calculations indicate that the conversion of 5 x 109 to 1.8 x 109 megagrams of carbon to the atmosphere.

Reconciling carbon-cycle concepts, terminology, and methods

Recent projections of climatic change have focused a great deal of scientific and public attention on patterns of carbon (C) cycling as well as its controls, particularly the factors that determine whether an ecosystem is a net source or sink of atmospheric carbon dioxide (CO2). Net ecosystem production (NEP), a central concept in C-cycling research, has been used by scientists to represent two different concepts. We propose that NEP be restricted to just one of its two original definitions—the imbalance between gross primary production (GPP) and ecosystem respiration (ER). We further propose that a new term—net ecosystem carbon balance (NECB)—be applied to the net rate of C accumulation in (or loss from [negative sign]) ecosystems. Net ecosystem carbon balance differs from NEP when C fluxes other than C fixation and respiration occur, or when inorganic C enters or leaves in dissolved form. These fluxes include the leaching loss or lateral transfer of C from the ecosystem; the emission of volatile organic C, methane, and carbon monoxide; and the release of soot and CO2 from fire. Carbon fluxes in addition to NEP are particularly important determinants of NECB over long time scales. However, even over short time scales, they are important in ecosystems such as streams, estuaries, wetlands, and cities. Recent technological advances have led to a diversity of approaches to the measurement of C fluxes at different temporal and spatial scales. These approaches frequently capture different components of NEP or NECB and can therefore be compared across scales only by carefully specifying the fluxes included in the measurements. By explicitly identifying the fluxes that comprise NECB and other components of the C cycle, such as net ecosystem exchange (NEE) and net biome production (NBP), we can provide a less ambiguous framework for understanding and communicating recent changes in the global C cycle.

A Carbon Budget for Forests of the Conterminous United States

The potential need for national-level comparisons of greenhouse gas emis- sions, and the desirability of understanding terrestrial sources and sinks of carbon, has prompted interest in quantifying national forest carbon budgets. In this study, we link a forest inventory database, a set of stand-level carbon budgets, and information on harvest levels in order to estimate the current pools and flux of carbon in forests of the conterminous United States. The forest inventory specifies the region, forest type, age class, productivity class, management intensity, and ownership of all timberland. The stand-level carbon bud- gets are based on growth and yield tables, in combination with additional information on carbon in soils, the forest floor, woody debris, and the understory. Total carbon in forests of the conterminous U.S. is estimated at 36.7 Pg, with half of that in the soil compartment. Tree carbon represents 33% of the total, followed by woody debris (10%), the forest floor (6%), and the understory (1%). The carbon uptake associated with net annual growth is 331 Tg, however, much of that is balanced by harvest-related mortality (266 Tg) and decomposition of woody debris. The forest land base at the national level is accumulating 79 Tg/yr, with the largest carbon gain in the Northeast region. The similarity in the mag- nitude of the biologically driven flux and the harvest-related flux indicates the importance of employing an age-class-based inventory, and of including effects associated with forest harvest and harvest residue, when modeling national carbon budgets in the temperate zone.

Tree Seedlings on Logs in Picea-Tsuga Forests of Oregon and Washington

Logs are the major seedbed for trees in coastal Picea sitchensis–Tsuga heterophylla forests. Field experiments were conducted at Cascade Head, Oregon, and Hoh River, Washington, to examine pathogens, predation, competition, and standing water as causes for this close seedling–log association. More seedlings survived on log blocks than on soil blocks, regardless of whether the blocks were raised or placed flush with the soil surface. Standing water was therefore an unlikely cause of the seedling–log association. Comparisons of plots protected from and exposed to predation revealed that predation was minor and of equal intensity on soils and logs. Sterilizing soils did not consistently increase seedling survival above controls. Clearing ground–layer vegetation from soil plots significantly increased the survival of conifer seedlings compared with that on uncleared soils. The seed penetration rates through moss mats indicated that <1% of the seedlings germinated within moss mats. Competition with herbs and mosses on the forest floor therefore appears to be responsible for the disproportionate number of tree seedlings found on logs. Recently fallen logs represent sites where competition is low enough for tree seedling recruitment within many Picea–Tsuga forests.

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.

A synthesis of current knowledge on forests and carbon storage in the United States

Using forests to mitigate climate change has gained much interest in science and policy discussions. We examine the evidence for carbon benefits, environmental and monetary costs, risks and trade-offs for a variety of activities in three general strategies: (1) land use change to increase forest area (afforestation) and avoid deforestation; (2) carbon management in existing forests; and (3) the use of wood as biomass energy, in place of other building materials, or in wood products for carbon storage. We found that many strategies can increase forest sector carbon mitigation above the current 162-256 Tg C/yr, and that many strategies have co-benefits such as biodiversity, water, and economic opportunities. Each strategy also has trade-offs, risks, and uncertainties including possible leakage, permanence, disturbances, and climate change effects. Because approximately 60% of the carbon lost through deforestation and harvesting from 1700 to 1935 has not yet been recovered and because some strategies store carbon in forest products or use biomass energy, the biological potential for forest sector carbon mitigation is large. Several studies suggest that using these strategies could offset as much as 10-20% of current U.S. fossil fuel emissions. To obtain such large offsets in the United States would require a combination of afforesting up to one-third of cropland or pastureland, using the equivalent of about one-half of the gross annual forest growth for biomass energy, or implementing more intensive management to increase forest growth on one-third of forestland. Such large offsets would require substantial trade-offs, such as lower agricultural production and non-carbon ecosystem services from forests. The effectiveness of activities could be diluted by negative leakage effects and increasing disturbance regimes. Because forest carbon loss contributes to increasing climate risk and because climate change may impede regeneration following disturbance, avoiding deforestation and promoting regeneration after disturbance should receive high priority as policy considerations. Policies to encourage programs or projects that influence forest carbon sequestration and offset fossil fuel emissions should also consider major items such as leakage, the cyclical nature of forest growth and regrowth, and the extensive demand for and movement of forest products globally, and other greenhouse gas effects, such as methane and nitrous oxide emissions, and recognize other environmental benefits of forests, such as biodiversity, nutrient management, and watershed protection. Activities that contribute to helping forests adapt to the effects of climate change, and which also complement forest carbon storage strategies, would be prudent.