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Conceptual 1 , 2 , 3 , 4 and numerical 5 , 6 , 7 , 8 models of nitrogen cycling in temperate forests assume that nitrogen is lost from these ecosystems predominantly by way of inorganic forms, such as nitrate and ammonium ions. Of these, nitrate is thought to be particularly mobile, being responsible for nitrogen loss to deep soil and stream waters. But human activities—such as fossil fuel combustion, fertilizer production and land-use change—have substantially altered the nitrogen cycle over large regions 9 , making it difficult to separate natural aspects of nitrogen cycling from those induced by human perturbations 10 . Here we report stream chemistry data from 100 unpolluted primary forests in temperate South America. Although the sites exhibit a broad range of environmental factors that influence ecosystem nutrient cycles 11 , 12 , 13 (such as climate, parent material, time of ecosystem development, topography and biotic diversity), we observed a remarkably consistent pattern of nitrogen loss across all forests. In contrast to findings from forests in polluted regions, streamwater nitrate concentrations are exceedingly low, such that nitrate to ammonium ratios were less than unity, and dissolved organic nitrogen is responsible for the majority of nitrogen losses from these forests. We therefore suggest that organic nitrogen losses should be considered in models of forest nutrient cycling, which could help to explain observations of nutrient limitation in temperate forest ecosystems.
Background China has committed to achieving peak CO 2 emissions before 2030 and carbon neutrality before 2060; therefore, accelerated efforts are needed to better understand carbon accounting in industry and energy fields as well as terrestrial ecosystems. The carbon sink capacity of plantation forests contributes to the mitigation of climate change. Plantation forests throughout the world are intensively managed, and there is an urgent need to evaluate the effects of such management on long-term carbon dynamics. Methods We assessed the carbon cycling patterns of ecosystems characterized by three typical plantation species (Chinese fir ( Cunninghamia lanceolata (Lamb.) Hook.), oak ( Cyclobalanopsis glauca (Thunb.) Oerst.), and pine ( Pinus massoniana Lamb.)) in Lishui, southern China, by using an integrated biosphere simulator (IBIS) tuned with localized parameters. Then, we used the state-and-transition simulation model (STSM) to study the effects of active forest management (AFM) on carbon storage by combining forest disturbance history and carbon cycle regimes. Results 1) The carbon stock of the oak plantation was lower at an early age (<50 years) but higher at an advanced age (>50 years) than that of the Chinese fir and pine plantations. 2) The carbon densities of the pine and Chinese fir plantations peaked at 70 years (223.36 Mg·ha ‒1 ) and 64 years (232.04 Mg·ha ‒1 ), respectively, while the carbon density in the oak plantation continued increasing (>100 years). 3) From 1989 to 2019, the total carbon pools of the three plantation ecosystems followed an upward trend (an annual increase of 0.16–0.22 Tg C), with the largest proportional increase in the aboveground biomass carbon pool. 4) AFM increased the recovery of carbon storage after 1996 and 2009 in the pine and Chinese fir plantations, respectively, but did not result in higher growth in the oak plantation. 5) The proposed harvest planning is reasonable and conducive to maximizing the carbon sequestration capacity of the forest. Conclusions This study provides an example of a carbon cycle coupling model that is potentially suitable for simulating China's plantation forest ecosystems and supporting carbon accounting to monitor peak CO 2 emissions and reach carbon neutrality.
We recognize three levels of extinction–global, local, and ecological – and provide examples of each. The protection and recovery of sea otters (Enhydra lutris) has provided abundant evidence of the consequences of their local extinction from kelp forest communities in the North Pacifc Ocean. These consequences include release of benthic invertebrate populations from limitation by predation; deforestation of kelp beds due to increased grazing by herbivorous sea urchins, one of the otter's main prey; and various cascading effects resulting from the biological and physical importance of kelp in coastal ecosystems. These interactions probably were important agents of selection for certain species. Two other examples are discussed: Steller's sea cow (Hydrodamalis gigas), a case of global extinction, and spiny lobsters, a possible case of ecological extinction. We speculate that grazing by sea cows was an important disturbance to surface-canopy-forming kelps and other algae in the littoral zones, but also point out that any such interactions probably acted in concert with physical disturbances by ocean waves. The ecological and evolutionary importance of sea cow grazing probably will remain a matter of speculation and conjecture because the species is globally extinct. Predation by spiny lobsters limits a variety of littoral and sublittoral invertebrate populations, particularly mollusks In one remarkable example, the reduction or local extinction of spiny lobsters enabled predutory whelks to increase in size and abundance, ultimately resulting in a predutor-prey role reversal. From these and other case studies we can clued that (1) the extinction of consumers may have brad and sometimes unexpected influences on kelp forest ecosystems; (2) direct or indirect interactions with now-extinct species probably exerted important selective influences on many extant forms; (3) such ecological and evolutionary influences are best understood where local or ecological extinctions, followed by recoveries, have provided comparisons in space or time; and (4) because of various ecological and behavioral barriers, local extinctions and their ecological consequences may not be simply reversed by protecting or reintroducing depleted or locally extinct species.
Acid deposition, more commonly known as acid rain, occurs when emissions of sulfur dioxide (SO2) and nitrogen oxides (NOx) react in the atmosphere (with water, oxygen, and oxidants) to form various acidic compounds. These acidic compounds then fall to earth in either a wet form (rain, snow, and fog) or a dry form (gases, aerosols, and particles). Prevailing winds transport the acidic compounds hundreds of miles, often across state and national borders. At certain levels the acidic compounds, including small particles such as sulfates and nitrates can cause many negative human health and environmental effects. While ecosystems are subject to many stresses, including land-use changes, climate change, and variations in hydrologic and meteorologic cycles, the scientific literature has clearly demonstrated that these pollutants can: • Degrade air quality, • Impair visibility, • Damage public health, • Acidify lakes and streams, • Harm sensitive forests, • Harm sensitive coastal ecosystems, and • Accelerate the decay of building materials, paints, and cultural artifacts such as buildings, statues, and sculptures. Title IV was passed by Congress as part of the 1990 Clean Air Act Amendments to reduce emissions of SO 2 and NO x from fossil fuel-burning power plants in order to protect ecosystems suffering damage from acid deposition and to improve air quality. At the same time, the National Acid Precipitation Assessment Program (NAPAP) was asked to periodically assess and report to Congress on the implementation of the Acid Rain Program, recent scientific knowledge surrounding acid deposition and its effects, and the reduction in acid deposition necessary to prevent adverse ecological effects. This NAPAP Report focuses primarily on emission reductions from power plants, both in terms of assessing past reductions under the Acid Rain Program and in projecting the ecological effects of additional reductions of SO 2 and NO x . It should be noted that power generation currently contributes approximately 69% of the SO 2 emissions and 22% of the NO x emissions nationwide. This contribution is decreasing as emissions from power generation continue to decrease, making the other sources of these pollutants more prominent. Modeling suggests that even if SO 2 emissions from power generation were reduced to zero, some lakes and streams would remain acidic due to acid deposition. However, there are several other regulations that reduce emissions of SO 2 and NO x from these non-power generation sources, such as the Tier II mobile source standards, the Heavy Duty Diesel standards, and the Clean Air Non-Road Diesel Rule, that have also been promulgated since 1990. These regulations, primarily designed to bring counties into attainment with fine particle and ozone air quality standards, also incidentally reduce emissions that contribute to acid deposition. Implementation of Title IV has successfully and substantially reduced emissions of SO 2 and NO x from power generation at a significantly lower cost than expected: • In 2002, SO 2 emissions were 10.2 million tons, 35% lower than 1990 emissions and 40% lower than 1980 emissions.* • In 2002, NO x emissions were 4.5 million tons, 33% lower than 1990 emissions. In addition, SO 2 emissions from all sources have decreased by 32% since 1990 and emissions of NO x from all sources have decreased by 12% since 1990. Power generating sources continue to close in on the goal of reducing power plant SO 2 emissions from 1980 levels by 50% (to 8.95 million tons) as required by the 1990 Clean Air Act. Power generating sources have also exceeded the goal of a two million ton reduction in NO x emissions from projected 2000 levels as required by the 1990 Clean Air Act. These emission reductions have contributed to measurable improvements in air quality, reductions in acid deposition, and the beginnings of recovery of acid-sensitive waters in some areas: • SO2 concentrations in the atmosphere (a precursor to fine particles and acid deposition) have decreased since 1990. Average annual SO2 concentrations in the Northeast in 2000–2002 were 40% lower than they were in 1989–1991, concentrations in the mid-Atlantic were 30% lower, concentrations in the Southeast were 35% lower, and concentrations in the Midwest were 45% lower. • Sulfate concentrations in the atmosphere (a major component of fine particles, especially in the East) have decreased since 1990 as well. Average annual sulfate concentrations in the Northeast and Midwest in 2000–2002 were approximately 30% lower than they were in 1989–1991, and concentrations in the mid-Atlantic and Southeast were 25% lower. • Wet sulfate deposition, a major component of acid rain, has also decreased since 1990. Average annual sulfate deposition in the Northeast in 2000–2002 was 40% lower than it was in 1989–1991, deposition in the mid-Atlantic and Midwest was 35% lower, and deposition in the Southeast was 25% lower. • Wet nitrate deposition has not decreased regionally from historical levels because of the relatively moderate NO x reduction from power plants and the continuing large contribution (over 50% of total NOx emissions) from other sources of NO x such as vehicles and nonroad vehicles. • Although visibility has begun to improve in some parts of the U.S., there is still significant impairment of visibility in many national parks and other Class I areas throughout the U.S. • Acid neutralizing capacity is beginning to rise in some surface waters in the Northeast, including lakes in the Adirondack Mountains (see graphic below). This is an indication that recovery from acidification is occurring in those areas.
Executive Summary The continued pressures of population growth on the life-sustaining, economic, and cultural ecosystem services provided by our national forests, particularly those located near rapidly growing urban areas, present ongoing challenges to forest managers. Achieving an effective assessment of these ecosystem services includes a proper accounting of the ecological, economic, and social values attributable to them. However, assessments of ecosystem goods and services notably lack information describing the spatial distribution and relative intensity of social values—the perceived, nonmarket values derived particularly from cultural ecosystem services. A geographic information system (GIS) tool developed to fill this need, Social Values for Ecosystem Services (SolVES; http://solves.cr.usgs.gov ), now provides the capability to generate social-value maps at a range of spatial scales. This report presents some of the methods behind SolVES, procedures needed to apply the tool, the first formal map products resulting from its application at a regional scale, and a discussion of the management implications associated with this type of information. In this study, we use SolVES to identify the location and relative intensity of social values as derived from survey responses gathered from residents living in counties adjacent to Arapaho, Roosevelt, Medicine Bow, Routt, and White River National Forests. The results, presented as a series of social-value maps, represent the first publicly available spatial data on social-value intensity for the southern Rocky Mountain region. Our analysis identified high-value areas for social values including aesthetic, biodiversity, and life sustaining within wilderness areas. Other values, like recreation, show high-value areas both within wilderness and throughout the general forest areas, which can be attributed to people using the forests for a diverse set of recreational activities. The economic social-value type was lower within wilderness areas, which was an expected outcome because of the restrictions inside wilderness areas that preclude resource extraction, development, and motorized or mechanized recreation. Providing spatially explicit social-value information collected from residents in counties adjacent to these national forests can assist in facilitating the U.S. Department of Agriculture Forest Service’s Multiple-Use Sustained-Yield Act of 1960 (16 U.S.C. 528) note and the 2012 Forest Service Planning Rule (36 CFR Part 219), which encourage public participation in planning and management. By making these maps available to the public, we hope to encourage exploration of potential uses of these data for resource management and planning.
Justifying the maintenance of small natural areas requires understanding their contribution to the conservation of specific natural resources. Mount Rushmore National Memorial (MORU) is a small portion of the Black Hills of western South Dakota and eastern Wyoming. Because it has been protected from logging since the late 1930s, it may serve as an important part of the Black Hills forest as a whole. To understand this role, we investigated the extent and degree of logging activities in the memorial and compared the current structure of the MORU forest to that in the rest of the Black Hills today and before Euro-American settlement. Our results suggest that approximately 29% of the park has had no tree harvesting activity, 18% of the park has had only selective cutting of trees, and 66% (344 ha) of the park's area is covered by old-growth forest. Based on current estimates of similar forest in the remainder of the Black Hills, the forest at MORU constitutes the second-largest area of old-growth ponderosa pine forest in the Black Hills. Although the current structure of the forest does not appear to be outside the range of natural variability for this ecosystem, some components of the forest are near the edge of this range. Conservation of this important natural resource will most likely require more active management than has occurred in the past, but this management will require careful consideration because of the rarity of this resource in the region.
The Moquah Barrens Research Natural Area (RNA) was established by the Chequamegon National Forest and the Lakes States Forest Experiment Station in 1935 with a research objective well-suited to the needs of the Forest Service and the scientific understanding of ecosystem function prevalent at the time of establishment. The original research plan was never implemented, which led to a joint Forest-Station decision in 1956 to disestablish the RNA. However, that decision was never implemented. A series of management decisions made after 1956 led to the loss of the pine barrens ecosystem originally encompassed by the RNA. This loss is not irretrievable and the work necessary to recover the original ecosystem is possible under existing RNA management guidelines. The experience of the Moquah Barrens RNA can be used by the Forest Service to improve overall management of the entire system of research natural areas. Two main areas of opportunity are identified: 1) implement an improved approach to managing official records associated with RNAs; and 2) adopt a management framework suitable for long-term ecological projects.
Sea level rise is leading to the rapid migration of marshes into coastal forests and other terrestrial ecosystems. Although complex biophysical interactions likely govern these ecosystem transitions, projections of sea level driven land conversion commonly rely on a simplified “threshold elevation” that represents the elevation of the marsh-upland boundary based on tidal datums alone. To determine the influence of biophysical drivers on threshold elevations, and their implication for land conversion, we examined almost 100,000 high-resolution marsh-forest boundary elevation points, determined independently from tidal datums, alongside hydrologic, ecologic, and geomorphic data in the Chesapeake Bay, the largest estuary in the U.S. located along the mid-Atlantic coast. We find five-fold variations in threshold elevation across the entire estuary, driven not only by tidal range, but also salinity and slope. However, more than half of the variability is unexplained by these variables, which we attribute largely to uncaptured local factors including groundwater discharge, microtopography, and anthropogenic impacts. In the Chesapeake Bay, observed threshold elevations deviate from predicted elevations used to determine sea level driven land conversion by as much as the amount of projected regional sea level rise by 2050. These results suggest that local drivers strongly mediate coastal ecosystem transitions, and that predictions based on elevation and tidal datums alone may misrepresent future land conversion.
The U.S. Geological Survey (USGS) was one of numerous governmental, private, and academic entities that provided input to the report The State of the Nation?s Ecosystems published periodically by the Heinz Center. This report describes the sources of data and methods used by the USGS to develop selected water?quality indicators for the 2007 edition of the Heinz Center report and documents modifications in the data sources and interpretations between the 2002 and 2007 editions of the Heinz Center report. Stream and ground?water quality data collected nationally as part of the USGS National Water-Quality Assessment Program were used to develop the ecosystem indicators for the Heinz Center report, including Core National indicators for the Movement of Nitrogen and Chemical Contamination and for selected ecosystems classified as Farmlands, Forest, Grasslands and Shrublands, Freshwater, and Urban and Suburban. In addition, the USGS provided water?quality and streamflow data collected as part of the National Stream Water Quality Accounting Network and the Federal?State Cooperative Program. The documentation provided herein serves not only as a reference for current and future editions of The State of the Nation?s Ecosystems but also provides critical information for future assessments of changes in contaminant occurrence in streams and ground water of the United States.
Restoration of tropical forests can lead to enhanced ecosystem services and increases in native biodiversity. Bryophytes may be an integral part of the forest restoration process and can serve a critical role in forest functioning. However, the recovery of bryophytes and their ability to facilitate woody plant establishment during restoration remains poorly studied, especially in the tropics. We investigate how bryophyte abundance and community composition, as well as woody plant seedling associations with bryophyte mats and other ground cover types change from under the canopy of intact forest to under trees in restoration corridor plantings in Hawaii. Restoration corridors consisted of corridors of koa (Acacia koa) trees that were planted roughly 30 years ago. Some corridors were planted around remnant ʻōhiʻa trees (Metrosideros polymorpha) that can be several hundred years old. We sampled under ʻōhiʻa in intact forest and both koa and ʻōhiʻa trees in restoration corridors. In restoration corridors, bryophyte abundance was low relative to intact forest and species diversity was a subset of that found in intact forest despite restoration corridors being several decades old. Seedlings strongly associated with bryophytes across all habitats suggesting that bryophytes may significantly enhance forest seedling establishment when present in restoration corridors. Other ground cover types like woody litter and nurse logs also had a positive association with forest seedlings but were rare in restoration corridors. Grass remained a dominant ground cover type in restoration corridors under koa and remnant ʻōhiʻa trees and only a single seedling was ever found growing in this grass. Enhancing bryophyte growth and recovery within restoration plantings through the reduction of grass cover could facilitate native plant establishment.
Introduction The mangrove forests across the Federated States of Micronesia provide critical resources and contribute to climate resilience. Locally, mangrove forests provide habitat for fish and wildlife, timber, and other cultural resources. Mangrove forests also protect Micronesian communities from tropical cyclones and tsunamis, providing a buffer against powerful waves and winds. Mangrove forests in Micronesia can store 700–1,800 metric tons of carbon per hectare (Donato and others, 2011), contributing to the estimated 5–10 billion metric tons of carbon stored by mangroves around the world (Alongi, 2018). This carbon storage is essential for global climate resilience. Mangrove forests and the benefits these ecosystems provide are threatened by accelerating sea-level rise and human activities. Healthy mangrove forests are resilient systems and have kept pace with some amounts of sea-level rise, but rapid sea-level rise could outpace the mangroves’ ability to adapt. Degraded mangroves are at greater risk where natural processes have been altered. Overharvest and clearing of timber, infrastructure development, and altered hydrology are just a few of the human activities that can damage mangrove forests.
Mangrove forests are highly productive ecosystems, but the fate of mangrove-derived carbon remains uncertain. Part of that uncertainty stems from the fact that gas transfer velocities in mangrove-surrounded waters are not well determined, leading to uncertainty in air-water CO 2 fluxes. Two SF 6 tracer release experiments were conducted to determine gas transfer velocities (k(600) = 8.3 ± 0.4 and 8.1 ± 0.6 cm h −1 ), along with simultaneous measurements of pCO 2 to determine the air-water CO 2 fluxes from Shark River, Florida (232.11 ± 23.69 and 171.13 ± 20.28 mmol C m −2 d −1 ), an estuary within the largest contiguous mangrove forest in North America. The gas transfer velocity results are consistent with turbulent kinetic energy dissipation measurements, indicating a higher rate of turbulence and gas exchange than predicted by commonly used wind speed/gas exchange parameterizations. The results have important implications for carbon fluxes in mangrove ecosystems.
Mangroves can increase their elevation relative to tidal flooding through biogeomorphic feedbacks but can submerge if rates of sea-level rise are too great. There is an urgent need to understand the vulnerability of mangroves to sea-level rise so local communities and resource managers can implement and prioritize actions. The need is especially pressing for small islands, which have been identified as an area of concern by the IPCC. We developed a generalizable modeling framework for tidal wetlands (WARMER-3) that accounts for species interactions and the belowground processes that dictate soil elevation building relative to sea levels. The model was calibrated with extensive field datasets, including accretion rates derived from 29 soil cores, over 300 forest inventory plots, water level, and elevation. The model included five mangrove tree species and was applied across seven regions around the Pacific Island of Pohnpei, Federated States of Micronesia, where mangrove forest is a critical ecosystem that supports subsistence living for local communities. We explored mangrove resilience and carbon accumulation under six sea-level rise scenarios. We also conducted an analysis to determine the sea-level rise rate threshold above which mangroves would be lost. The results suggest that Pohnpei mangroves can build their elevations relative to low and moderate rates of sea-level rise to prevent submergence, with limited loss of mangrove area through 2150. Under higher sea-level rise rates, however, forest elevation decreased substantially relative to mean sea level and there was extensive loss of mangrove area by that year. Regarding mangrove community composition, for all sea-level rise scenarios, the model predicted a change to increasing relative abundance of flood tolerant species and decreasing relative abundance of high-elevation species, which started to being realized by 2100. Variation in sediment supply, water levels, and elevation capital led to differential vulnerability around the island. We identified a threshold for Pohnpei mangroves where if local sea-level rise rates exceed 7.8 ± 2.2 mm/year they are projected to eventually submerge and be lost. Our modeling framework is novel by addressing both species interactions and critical belowground processes to better understand potential tidal ecosystem responses to sea-level rise.
Acidic deposition is the transfer of strong acids and acid-forming substances from the atmosphere to the surface of the Earth. The composition of acidic deposition includes ions, gases, and particles derived from the following: gaseous emissions of sulfur dioxide (SO 2 ), nitrogen oxides (NO x ), ammonia (NH 3 ), and particulate emissions of acidifying and neutralizing compounds. Over the past quarter century of study, acidic deposition has emerged as a critical environmental stress that affects forested landscapes and aquatic ecosystems in North America, Europe, and Asia. This complex problem is an example of a new class of environmental issues that are multiregional in scale and are not amenable to simple resolution by policymakers. Acidic deposition can originate from transboundary air pollution and can affect large geographic areas. It is highly variable across space and time, links air pollution to diverse terrestrial and aquatic ecosystems, and alters the interactions of many elements (e.g., sulfur [S], nitrogen [N], hydrogen ion [H + ], calcium [Ca 2+ ], magnesium [Mg 2+ ], and aluminum [Al]). It also contributes directly and indirectly to biological stress and to the degradation of ecosystems. Despite the complexity of the effects of acidic deposition, North American and European management actions directed toward the recovery of damaged natural resources have resulted in recent decreases in both emissions and deposition of acidic S compounds. Accordingly, acidic deposition presents an instructive case study for coordinating science and policy efforts aimed at resolving large-scale environmental problems.
Chemistry of precipitation and streamwater and resulting input-output budgets for nutrient ions were determined concurrently for three years on three upland, forested watersheds located within an 80 km radius in central New England. Chemistry of precipitation and inputs of nutrients via wet deposition were similar among the three watersheds and were generally typical of central New England. In contrast, chemistry and nutrient outputs in streamwater varied dramatically between watersheds, with chemistries ranging from acidic to alkaline. Comparisons with data reported for 159 other upland, forested watersheds in central New England show that our study watersheds span the regional range likely to be encountered in stream chemistry. The regional variability stems in part from past natural disturbances such as wildfire, and variations in source of soil parent material. An approach is presented for predicting the important influence of glacial till on stream chemistry, including acid-base relationships, aluminum content, and nutrient outputs. Knowledge of streamwater chemistry and controlling factors can serve as an index of how terrestrial and aquatic ecosystems will respond to forest management activities and atmospheric deposition.
Evapotranspiration (ET) is the largest component of water loss from terrestrial ecosystems; however, large uncertainties exist when estimating the temporal and spatial variations of ET because of concurrent shifts in the magnitude and seasonal distribution of precipitation as well as differences in the response of ecosystem ET to environmental variabilities. In this study, we examined the impacts of precipitation seasonality and ecosystem types on ET quantified by eddy covariance towers from 2002 to 2004 in three ecosystems (grassland, deciduous broadleaf forest, and evergreen needleleaf forest) in the Yukon River Basin, Alaska. The annual precipitation changed greatly in both magnitude and seasonal distribution through the three investigated years. Observations and model results showed that ET was more sensitive to precipitation scarcity in the early growing season than in the late growing season, which was the direct result of different responses of ET components to precipitation in different seasons. The results demonstrated the importance of seasonal variations of precipitation in regulating annual ET and overshadowing the function of annual precipitation. Comparison of ET among ecosystems over the growing season indicated that ET was largest in deciduous broadleaf, intermediate in evergreen needleleaf, and lowest in the grassland ecosystem. These ecosystem differences in ET were related to differences in successional stages and physiological responses.
The decline of Taxodium distichum forests along the Gulf Coast of North America is partly due to elevation loss and subsequent flooding. In many coastal wetlands, a common approach for coastal restoration is to rebuild elevation through the application of dredge material, but this technique has not been used widely in coastal forests due to concerns of negatively impacting trees. This experiment explored growth responses of Nyssa aquatica and T. distichum saplings to applications of low salinity dredge material (0.08 ± 0.001 ppt) in a greenhouse setting. Compared to controls, saplings of T. distichum grown in 7 and 15 cm sediment depths had greater final height, and increased stem and total biomass. In contrast, N. aquatica did not respond to sediment application. The absence of a negative response to sediment application in these two species indicates that dredge material application has the potential to improve the ecosystem health of sinking swamp forests by raising their elevation. We recommend that field trials applying sediment additions in coastal forests include careful monitoring of ecosystem responses, including seed bank expression, seedling regeneration, and root and canopy production.