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Society is confronted by interconnected threats to ecological sustainability. Among these is the devastation of forests by destructive non-native pathogens and insects introduced through global trade, leading to the loss of critical ecosystem services and a global forest health crisis. We argue that the forest health crisis is a public-good social dilemma and propose a response framework that incorporates principles of collective action. This framework enables scientists to better engage policymakers and empowers the public to advocate for proactive biosecurity and forest health management. Collective action in forest health features broadly inclusive stakeholder engagement to build trust and set goals; accountability for destructive pest introductions; pooled support for weakest-link partners; and inclusion of intrinsic and nonmarket values of forest ecosystems in risk assessment. We provide short-term and longer-term measures that incorporate the above principles to shift the societal and ecological forest health paradigm to a more resilient state.

Although awareness that air pollution can damage vegetation dates back at least to the 1600s, the processes and mechanisms of damage were not rigorously studied until the late twentieth century. In the UK following the Industrial Revolution, urban air quality became very poor, with highly phytotoxic SO 2 and NO 2 concentrations, and remained that way until the mid-twentieth century. Since then both air quality, and our understanding of pollutants and their impacts, have greatly improved. Air pollutants remain a threat to natural and managed ecosystems. Air pollution imparts impacts through four major threats to vegetation are discussed through in a series of case studies. Gas-phase effects by the primary emissions of SO 2 and NO 2 are discussed in the context of impacts on lichens in urban areas. The effects of wet and dry deposited acidity from sulfur and nitrogen compounds are considered with a particular focus on forest decline. Ecosystem eutrophication by nitrogen deposition focuses on heathland decline in the Netherlands, and ground-level ozone at phytotoxic concentrations is discussed by considering impacts on semi-natural vegetation. We find that, although air is getting cleaner, there is much room for additional improvement, especially for the effects of eutrophication on managed and natural ecosystems. This article is part of a discussion meeting issue 'Air quality, past present and future'.

Little is known about the regional extent and variability of nitrate from atmospheric deposition that is transported to streams without biological processing in forests. We measured water chemistry and isotopic tracers (δ 18 O and δ 15 N) of nitrate sources across the Northern Forest Region of the U.S. and Canada and reanalyzed data from other studies to determine when, where, and how unprocessed atmospheric nitrate was transported in catchments. These inputs were more widespread and numerous than commonly recognized, but with high spatial and temporal variability. Only 6 of 32 streams had high fractions (>20%) of unprocessed atmospheric nitrate during baseflow. Seventeen had high fractions during stormflow or snowmelt, which corresponded to large fractions in near-surface soil waters or groundwaters, but not deep groundwater. The remaining 10 streams occasionally had some (<20%) unprocessed atmospheric nitrate during stormflow or baseflow. Large, sporadic events may continue to be cryptic due to atmospheric deposition variation among storms and a near complete lack of monitoring for these events. A general lack of observance may bias perceptions of occurrence; sustained monitoring of chronic nitrogen pollution effects on forests with nitrate source apportionments may offer insights needed to advance the science as well as assess regulatory and management schemes.

Historical increases in emissions and atmospheric deposition of oxidized and reduced nitrogen (N) provided the impetus for extensive, global-scale research investigating the effects of excess N in terrestrial and aquatic ecosystems, with several regions within the Eastern Deciduous Forest of the United States found to be susceptible to negative effects of excess N. The Clean Air Act and associated rules have led to decreases in emissions and deposition of oxidized N, especially in eastern U.S., representing a research challenge and opportunity for ecosystem ecologists and biogeochemists. The purpose of this paper is to predict changes in the structure and function of North American forest ecosystems in a future of decreased N deposition. Hysteresis is a property of a system wherein output is not a strict function of corresponding input, incorporating lag, delay, or history dependence, particularly when the response to decreasing input is different from the response to increasing input. We suggest a conceptual hysteretic model predicting varying lag times in recovery of soil acidification, plant biodiversity, soil microbial communities, forest carbon (C) and N cycling, and surface water chemistry toward pre-N impact conditions. Nearly all of these can potentially respond strongly to reductions in N deposition. Most responses are expected to show some degree of hysteresis, with the greatest delays in response occurring in processes most tightly linked to "slow pools" of N in wood and soil organic matter. Because experimental studies of declines in N loads in forests of North America are lacking and because of the expected hysteresis, it is difficult to generalize from experimental results to patterns expected from declining N deposition. These will likely be long-term phenomena, difficult to distinguish from other, concurrent environmental changes, including elevated atmospheric CO 2 , climate change, reductions in acidity, invasions of new species, and long-term vegetation responses to past disturbance., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

We review and synthesize information on invasions of nonnative forest insects and diseases in the United States, including their ecological and economic impacts, pathways of arrival, distribution within the United States, and policy options for reducing future invasions. Nonnative insects have accumulated in United States forests at a rate of ~2.5 per yr over the last 150 yr. Currently the two major pathways of introduction are importation of live plants and wood packing material such as pallets and crates. Introduced insects and diseases occur in forests and cities throughout the United States, and the problem is particularly severe in the Northeast and Upper Midwest. Nonnative forest pests are the only disturbance agent that has effectively eliminated entire tree species or genera from United States forests within decades. The resulting shift in forest structure and species composition alters ecosystem functions such as productivity, nutrient cycling, and wildlife habitat. In urban and suburban areas, loss of trees from streets, yards, and parks affects aesthetics, property values, shading, stormwater runoff, and human health. The economic damage from nonnative pests is not yet fully known, but is likely in the billions of dollars per year, with the majority of this economic burden borne by municipalities and residential property owners. Current policies for preventing introductions are having positive effects but are insufficient to reduce the influx of pests in the face of burgeoning global trade. Options are available to strengthen the defenses against pest arrival and establishment, including measures taken in the exporting country prior to shipment, measures to ensure clean shipments of plants and wood products, inspections at ports of entry, and post-entry measures such as quarantines, surveillance, and eradication programs. Improved data collection procedures for inspections, greater data accessibility, and better reporting would support better evaluation of policy effectiveness. Lack of additional action places the nation, local municipalities, and property owners at high risk of further damaging and costly invasions. Adopting stronger policies to reduce establishments of new forest insects and diseases would shift the major costs of control to the source and alleviate the economic burden now borne by homeowners and municipalities., (© 2016 by the Ecological Society of America.)

Climate change and atmospheric nitrogen (N) deposition are two of the most important global change drivers. However, the interactions of these drivers have not been well studied. We aimed to assess how the combined effect of soil N additions and more frequent soil drying-rewetting events affects carbon (C) and N cycling, soil:atmosphere greenhouse gas (GHG) exchange, and functional microbial diversity. We manipulated the frequency of soil drying-rewetting events in soils from ambient and N-treated plots in a temperate forest and calculated the Orwin & Wardle Resistance index to compare the response of the different treatments. Increases in drying-rewetting cycles led to reductions in soil NO3- levels, potential net nitrification rate, and soil : atmosphere GHG exchange, and increases in NH4+ and total soil inorganic N levels. N-treated soils were more resistant to changes in the frequency of drying-rewetting cycles, and this resistance was stronger for C- than for N-related variables. Both the long-term N addition and the drying-rewetting treatment altered the functionality of the soil microbial population and its functional diversity. Our results suggest that increasing the frequency of drying-rewetting cycles can affect the ability of soil to cycle C and N and soil : atmosphere GHG exchange and that the response to this increase is modulated by soil N enrichment., (© 2015 John Wiley & Sons Ltd.)

Repeated invasions of non-native insects and pathogens have altered the structure and function of forest ecosystems in the Catskill Mountains of New York State, and will continue to do so in the future. Gypsy moth, beech bark disease, and hemlock woolly adelgid are among the insects and diseases currently established in the Catskills that are having significant effects on forests. Many others, including emerald ash borer, Asian long-horned beetle, Phytophthora ramorum, and Sirex wood wasp, are either very recently established in the Catskills or have been found elsewhere in North America and threaten to spread to this region. Short-term disturbances associated with these pests include reduction of productivity, tree decline and mortality, disruption of nutrient cycles, and reduction of seed production. Longer-term impacts are associated with shifts in tree species composition that alter productivity, nutrient cycling, and biodiversity. Catskill forests at mid to high elevations, such as the New York State Forest Preserve lands, are dominated by sugar maple and are particularly vulnerable to pests that use maple as a host, including the Asian long-horned beetle. The simultaneous effects of multiple invading insects and pathogens, and their interactions with changing climate and air pollution regimes, make it very difficult to predict the future composition of Catskill forests., (© 2013 New York Academy of Sciences.)

Stress within tree roots may influence whole-tree responses to nutrient deficiencies or toxic ion accumulation, but the mechanisms that govern root responses to the belowground chemical environment are poorly quantified. Currently, root production is modeled using rates of forest production and stoichiometry, but this approach alone may be insufficient to forecast variability in forest responses when physical and chemical stressors alter root lifespan, rooting depth or mycorrhizal colonization directly. Here, we review key research priorities for improving predictions of tree responses to changes in the belowground biogeochemical environment resulting from nitrogen deposition, including: limits of the optimum allocation paradigm, root physiological stress and lifespan, contingency effects that determine threshold responses across broad gradients, coupled water-biogeochemical interactions on roots, mycorrhizal dynamics that mediate root resilience and model frameworks to better simulate root feedbacks to aboveground function. We conclude that models incorporating physiological feedbacks, dynamic responses to coupled stressors, mycorrhizal interactions, and which challenge widely-accepted notions of optimum allocation, can elucidate potential thresholds of tree responses to biogeochemical stressors. Emphasis on comparative studies across species and environmental gradients, and which incorporates insights at the cellular and ecosystem level, is critical for forecasting whole-tree responses to altered biogeochemical landscapes.

Climate change predictions for the northeastern US call for an increase in tropical storms and a decrease in extra tropical cyclones including continental storms. We ran 24-h back trajectories for each precipitation event that occurred at the Cary Institute of Ecosystem Studies in southeastern New York, U.S.A. from 1984 to 2007 and analyzed precipitation chemistry as well as air mass position 24 h prior to the onset of each precipitation event. The results showed an increase in marine precipitation and a slight but statistically insignificant decrease in continental precipitation during the 1984-2007 period. The chemistry of precipitation from the two directions was quite different marine storms were higher in Na4 and Cl- but lower in solutes associated with acid precipitation (H+, SO4(2-), NO3-, and NH4+). Annual mean concentrations of acid precipitation solutes declined for storms from both directions during the period. We used a simple mixing model based on the current rates of increase and decrease of marine and continental precipitation respectively to show that chemical changes in precipitation resulting from the shift in storm tracks are small compared to chemical changes due to emissions reductions.

Conservation organizations have most often focused on land-use change, climate change, and invasive species as prime threats to biodiversity conservation. Although air pollution is an acknowledged widespread problem, it is rarely considered in conservation planning or management. In this synthesis, the state of scientific knowledge on the effects of air pollution on plants and animals in the Northeastern and Mid-Atlantic regions of the United States is summarized. Four air pollutants (sulfur, nitrogen, ozone, and mercury) and eight ecosystem types ranging from estuaries to alpine tundra are considered. Effects of air pollution were identified, with varying levels of certainty, in all the ecosystem types examined. None of these ecosystem types is free of the impacts of air pollution, and most are affected by multiple pollutants. In aquatic ecosystems, effects of acidity, nitrogen, and mercury on organisms and biogeochemical processes are well documented. Air pollution causes or contributes to acidification of lakes, eutrophication of estuaries and coastal waters, and mercury bioaccumulation in aquatic food webs. In terrestrial ecosystems, the effects of air pollution on biogeochemical cycling are also very well documented, but the effects on most organisms and the interaction of air pollution with other stressors are less well understood. Nevertheless, there is strong evidence for effects of nitrogen deposition on plants in grasslands, alpine areas, and bogs, and for nitrogen effects on forest mycorrhizae. Soil acidification is widespread in forest ecosystems across the eastern United States and is likely to affect the composition and function of forests in acid-sensitive areas over the long term. Ozone is known to cause reductions in photosynthesis in many terrestrial plant species. For the most part, the effects of these pollutants are chronic, not acute, at the exposure levels common in the eastern United States. Mortality is often observed only at experimentally elevated exposure levels or in combination with other stresses such as drought, freezing, or pathogens. The notable exceptions are the acid/aluminum effects on aquatic organisms, which can be lethal at levels of acidity observed in many surface waters in the region. Although the effects are often subtle, they are important to biological conservation. Changes in species composition caused by terrestrial or aquatic acidification or eutrophication can propagate throughout the food webs to affect many organisms beyond those that are directly sensitive to the pollution. Likewise, sublethal doses of toxic pollutants may reduce the reproductive success of the affected organisms or make them more susceptible to potentially lethal pathogens. Many serious gaps in knowledge that warrant further research were identified. Among those gaps are the effects of acidification, ozone, and mercury on alpine systems, effects of nitrogen on species composition of forests, effects of mercury in terrestrial food webs, interactive effects of multiple pollutants, and interactions among air pollution and other environmental changes such as climate change and invasive species. These gaps in knowledge, coupled with the strong likelihood of impacts on ecosystems that have not been studied in the region, suggests that current knowledge underestimates the actual impact of air pollutants on biodiversity. Nonetheless, because known or likely impacts of air pollution on the biodiversity and function of natural ecosystems are widespread in the Northeast and Mid-Atlantic regions, the effects of air pollution should be considered in any long-term conservation strategy. It is recommended that ecologically relevant standards, such as "critical loads," be adopted for air pollutants and the importance of long-term monitoring of air pollution and its effects is emphasized.

Sodium and chloride concentrations and export increased from 1986 to 2005 in a rural stream in southeastern New York. Concentrations increased 1.5 mg/L per year (chloride) and 0.9 mg/L per year (sodium), and export increased 33,000 kg/year (chloride) and 20,000 kg/year (sodium) during this period. We estimate that salt used for deicing accounted for 91% of the sodium chloride input to the watershed, while sewage and water softeners accounted for less than 10% of the input. Road salt use in the watershed did not increase during the study, but sodium and chloride from sewage and water softeners is likely to have increased slightly due to a small increase in population. Increased input from sewage and water softeners cannot account for the increase in concentration and export from the watershed. Model results suggest that the increase in streamwater concentration and export was likely due to a lag effect of long-term road salt use and subsurface buildup.

Watersheds within the Catskill Mountains, New York, receive among the highest rates of nitrogen (N) deposition in the northeastern United States and are beginning to show signs of N saturation. Despite similar amounts of N deposition across watersheds within the Catskill Mountains, rates of soil N cycling and N retention vary significantly among stands of different tree species. We examined the potential use of delta (15)N of plants and soils as an indicator of relative forest soil N cycling rates. We analyzed the delta (15)N of foliage, litterfall, bole wood, surface litter layer, fine roots and organic soil from single-species stands of American beech (Fagus grandifolia), eastern hemlock (Tsuga canadensis), red oak (Quercus rubra), and sugar maple (Acer saccharum). Fine root and organic soil delta (15)N values were highest within sugar maple stands, which correlated significantly with higher rates of net mineralization and nitrification. Results from this study suggest that fine root and organic soil delta (15)N can be used as an indicator of relative rates of soil N cycling. Although not statistically significant, delta (15)N was highest within foliage, wood and litterfall of beech stands, a tree species associated with intermediate levels of soil N cycling rates and forest N retention. Our results show that belowground delta (15)N values are a better indicator of relative rates of soil N cycling than are aboveground delta (15)N values.

Atmospheric deposition has long been recognized as an important source of pollutants and nutrients to ecosystems. The need for reliable, spatially explicit estimates of total atmospheric deposition (wet + dry + cloud) is central, not only to air pollution effects researchers, but also for calculation of input-output budgets, and to decision makers faced with the challenge of assessing the efficacy of policy initiatives related to deposition. Although atmospheric deposition continues to represent a critical environmental and scientific issue, current estimates of total deposition have large uncertainties, particularly across heterogeneous landscapes such as montane regions. We developed an empirical modeling approach that predicts total deposition as a function of landscape features. We measured indices of total deposition to the landscapes of Acadia (121 km2) and Great Smoky Mountains (2074 km2) National Parks (USA). Using approximately 300-400 point measurements and corresponding landscape variables at each park, we constructed a statistical (general linear) model relating the deposition index to landscape variables measured in the field. The deposition indices ranged over an order of magnitude, and in response to vegetation type and elevation, which together explained approximately 40% of the variation in deposition. Then, using the independent landscape variables available in GIS data layers, we created a GIS-relevant statistical nitrogen (N) and sulfur (S) deposition model (LandMod). We applied this model to create park-wide maps of total deposition that were scaled to wet and dry deposition data from the closest national network monitoring stations. The resultant deposition maps showed high spatial heterogeneity and a four- to sixfold variation in "hot spots" and "cold spots" of N and S deposition ranging from 3 to 31 kg N x ha(-1) x yr(-1) and from 5 to 42 kg S x ha(-1) x yr(-1) across these park landscapes. Area-weighted deposition was found to be up to 70% greater than NADP plus CASTNET monitoring-station estimates together. Model-validation results suggest that the model slightly overestimates deposition for deciduous and coniferous forests at low elevation and underestimates deposition for high-elevation coniferous forests. The spatially explicit deposition estimates derived from LandMod are an improvement over what is currently available. Future research should test LandMod in other mountainous environments and refine it to account for (currently) unexplained variation in deposition.

Ammonium (NH(4)(+)) concentrations in air and precipitation at the Institute of Ecosystem Studies (IES) in southeastern New York, USA declined over an 11-year period from 1988 to 1999, but increased from 1999 to 2001. These trends in particulate NH(4)(+) correlated well with trends in particulate SO(4)(2-) over the 1988-2001 period. The NH(4)(+) trends were not as well correlated with local cattle and milk production, which declined continuously throughout the period. This suggests that regional transport of SO(4)(2-) may have a greater impact on concentrations of NH(4)(+) and subsequent deposition than local agricultural emissions of NH(3). Ammonium concentrations in precipitation correlated significantly with precipitation SO(4)(2-) concentrations for the 1984-2001 period although NH(4)(+) in precipitation increased after 1999 and SO(4)(2-) in precipitation continued to decline after 1999. The correlation between NH(4)(+) and SO(4)(2-) was stronger for particulates than for precipitation. Particulate NH(4)(+) concentrations were also correlated with particulate SO(4)(2-) concentrations at 31 of 35 eastern U.S. CASTNet sites that had at least 10 years of data. Air concentrations of NH(4)(+) and SO(4)(2-) were more strongly correlated at the sites that were located within an agricultural landscape than in forested sites. At most of the sites there was either no trend or a decrease in NH(4)(+) dry deposition during the 1988-2001 period. The sites that showed an increasing trend in NH(4)(+) dry deposition were generally located in the southeastern U.S. The results of this study suggest that, in the northeastern U.S., air concentrations of NH(4)(+) and subsequent deposition may be more closely linked to SO(4)(2-) and thus SO(2) emissions than with NH(3) emissions. These results also suggest that reductions in S emissions have reduced NH(4)(+) transport to and NH(4)(+)-N deposition in the Northeast.

Forest defoliation by insects can lead to severe disruptions of the nitrogen (N) cycle resulting in elevated NO 3 - levels in stream water. To trace the movement of insect-mobilized N in a forest soil, 15 N-labeled gypsy moth frass or 15 N-labeled oak leaf litter was added to trenched plots in an oak forest over 29 months. Nitrogen movement from the frass or litter was measured in the available, mineralizable, microbial and total soil pools. Uptake of 15 N by oak seedlings and inorganic N leaching losses were also measured. No significant differences were found between the frass or leaf treatments for total N in any of the pools. Significant differences were found among the treatments in the distribution of the 15 N tracer. Forty percent of the 15 N added as frass became incorporated in the soils, with less than 1% found in oak seedlings. Almost 80% of 15 N added as leaves remained in the undecomposed leaf material after 2 years. Less than 0.001% of the added 15 N was leached in both treatments. Our data indicate that N in frass is mobilized more quickly than N in leaf litter. However, this frass N may be largely unavailable to plants and microorganisms as little of it was found in the extractable, microbial, or readily mineralizable pools.

The gypsy moth (Lymantria dispar L.), a major defoliator of hardwood forests in the eastern U.S., has a highly alkaline midgut pH. We hypothesized that the high pH would cause high rates of ammonia (NH 3 ) volatilization as larvae consumed foliage, leading to potentially large losses of N from the ecosystem to the atmosphere during gypsy moth outbreaks. We measured NH 3 emission during the consumption of oak foliage by larvae in the laboratory. Surprisingly, we found very low amounts of NH 3 release of about 0.1% of the N consumed in foliage. We speculate that digestive mechanisms may limit NH 3 production in the midgut, and that the acidic environment of the hindgut traps most of the small amount of NH 3 that is produced, effectively preventing a potentially very large N loss from both larvae and ecosystem. The estimated rate of NH 3 emission from a defoliated forest is small compared to other inputs and outputs of N from the ecosystem, but could potentially enhance the neutralization of atmospheric acidity during the defoliation period.

Defoliation of forests by insects is often assumed to produce a pulse of available nitrogen (N) from the decomposition of frass pellets. In this study we measured rates of carbon (C) and N mineralization from gypsy moth frass incubated with and without soil, and for soil alone. Incubations were at constant temperature and soil moisture conditions and lasted for 120 days. We found that gypsy moth frass contains much labile C as well as extractable N, and that the stimulation of microbial growth by the labile C results in immobilization of essentially all of the extractable N in the frass. The response of the microbes is fast, beginning within 1 day and lasting at least 90 days. This immobilization response represents an efficient mechanism for conserving N within a forest ecosystem after a defoliation event.

Spatial patterns of atmospheric deposition across the northeastern United States were evaluated and summarized in a simple model as a function of elevation and geographic position within the region. For wet deposition, 3-11 yr of annual concentration data for the major ions in precipitation were obtained from the National Atmospheric Deposition Program/National Trend Network (NADP/NTN) for 26 sites within the region. Concentration trends were evaluated by regression of annual mean concentrations against latitude and longitude. For nitrate, sulfate, and ammonium concentrations, a more than twofold linear decrease occurs from western New York and Pennsylvania to eastern Maine. These trends were combined with regional and elevational trends of precipitation amount, obtained from 30-yr records of annual precipitation at >300 weather stations, to provide long-term patterns of wet deposition. Regional trends of dry deposition of N and S compounds were determined using 2-3 yr of particle and gas concentration data collected by the National Dry Deposition Network (NDDN) and several other sources, in combination with estimates of deposition velocities. Contrary to wet deposition trends, the dominant air concentration trends were steep decreases from south to north, creating regional decreases in total deposition (wet + dry) from the southwest to the northeast. This contrast between wet and dry deposition trends suggests that within the northeast the two deposition forms are received in different proportions from different source areas, wet deposited materials primarily from areas to the west and dry deposited materials primarily from urban areas along the southern edge of the region. The equations generated describing spatial patterns of wet and dry deposition within the region were entered into a geographic information system (GIS) containing a digital elevation model (DEM) in order to develop spatially explicit predictions of atmospheric deposition for the region., (© 1993 by the Ecological Society of America.)

To examine how rates of net photosynthesis and N uptake of red oak seedlings respond to defoliation under contrasting conditions of N availability, nitrogen-deficient plants were grown in sand culture and subjected to partial defoliation and increased N availability under low light conditions. Both photosynthesis and N uptake rates were measured regularly before and after the treatments. Defoliation resulted in elevated rates of net photosynthesis in both low-N and high-N trees, but the high-N trees were able to maintain the high photosynthetic rates for a longer period of time. Nitrogen availability did not affect the photosynthetic rate of the undefoliated plants. Nitrogen uptake was not affected by the defoliation treatment, but was increased by increasing N availability in both the defoliated and undefoliated plants. Nitrogen uptake rates increased less than would be expected on the basis of N availability alone, but the uptake rates were apparently not limited by carbon supply in the short term. Suboptimal concentrations of N in plant tissues resulted in a strong sink for N even in the absence of refoliation.

Airborne particles and vapors contributed significantly to the nutrient requirements and the pollutant load of a mixed hardwood forest in the eastern United States. Dry deposition was an important mechanism of atmospheric input to the foliar canopy, occurring primarily by vapor uptake for sulfur, nitrogen, and free acidity and by particle deposition for calcium and potassium. The canopy retained 50 to 70 percent of the deposited free acidity and nitrogen, but released calcium and potassium. Atmospheric deposition supplied 40 and 100 percent of the nitrogen and sulfur requirements, respectively, for the annual woody increment. This contribution was underestimated significantly by standard bulk deposition collectors.

Subalpine forests of the northern Appalachians are subject to significant deposition of water and chemicals via cloud droplet impaction. This deposition has been estimated by a method linking micrometeorological measures of turbulent transfer, a detailed representation of canopy structure, and experimentally derived capture efficiencies. Water inputs from clouds are about 46 percent, and chemical inputs range from 150 to 430 percent of the bulk precipitation.