Mark E. Fenn

Global assessment of nitrogen deposition effects on terrestrial plant diversity: a synthesis

Atmospheric nitrogen (N) deposition is a recognized threat to plant diversity in temperate and northern parts of Europe and North America. This paper assesses evidence from field experiments for N deposition effects and thresholds for terrestrial plant diversity protection across a latitudinal range of main categories of ecosystems, from arctic and boreal systems to tropical forests. Current thinking on the mechanisms of N deposition effects on plant diversity, the global distribution of G200 ecoregions, and current and future (2030) estimates of atmospheric N-deposition rates are then used to identify the risks to plant diversity in all major ecosystem types now and in the future. This synthesis paper clearly shows that N accumulation is the main driver of changes to species composition across the whole range of different ecosystem types by driving the competitive interactions that lead to composition change and/or making conditions unfavorable for some species. Other effects such as direct toxicity of nitrogen gases and aerosols, long-term negative effects of increased ammonium and ammonia availability, soil-mediated effects of acidification, and secondary stress and disturbance are more ecosystem- and site-specific and often play a supporting role. N deposition effects in mediterranean ecosystems have now been identified, leading to a first estimate of an effect threshold. Importantly, ecosystems thought of as not N limited, such as tropical and subtropical systems, may be more vulnerable in the regeneration phase, in situations where heterogeneity in N availability is reduced by atmospheric N deposition, on sandy soils, or in montane areas. Critical loads are effect thresholds for N deposition, and the critical load concept has helped European governments make progress toward reducing N loads on sensitive ecosystems. More needs to be done in Europe and North America, especially for the more sensitive ecosystem types, including several ecosystems of high conservation importance. The results of this assessment show that the vulnerable regions outside Europe and North America which have not received enough attention are ecoregions in eastern and southern Asia (China, India), an important part of the mediterranean ecoregion (California, southern Europe), and in the coming decades several subtropical and tropical parts of Latin America and Africa. Reductions in plant diversity by increased atmospheric N deposition may be more widespread than first thought, and more targeted studies are required in low background areas, especially in the G200 ecoregions.

Nitrogen excess in North American ecosystems: predisposing factors, ecosystem responses, and management strategies

Most forests in North America remain nitrogen limited, although recent studies have identified forested areas that exhibit symptoms of N excess, analogous to overfertilization of arable land. Nitrogen excess in watersheds is detrimental because of disruptions in plant/soil nutrient relations, increased soil acidification and aluminum mobility, increased emissions of nitrogenous greenhouse gases from soil, reduced methane consumption in soil, decreased water quality, toxic effects on freshwater biota, and eutrophication of coastal marine waters. Elevated nitrate (NO3−) loss to groundwater or surface waters is the primary symptom of N excess. Additional symptoms include increasing N concentrations and higher N:nutrient ratios in foliage (i.e., N:Mg, N:P), foliar accumulation of amino acids or NO3−, and low soil C:N ratios. Recent nitrogen-fertilization studies in New England and Europe provide preliminary evidence that some forests receiving chronic N inputs may decline in productivity and experience greate...

Ecological Effects of Nitrogen Deposition in the Western United States

Abstract In the western United States vast acreages of land are exposed to low levels of atmospheric nitrogen (N) deposition, with interspersed hotspots of elevated N deposition downwind of large, expanding metropolitan centers or large agricultural operations. Biological response studies in western North America demonstrate that some aquatic and terrestrial plant and microbial communities are significantly altered by N deposition. Greater plant productivity is counterbalanced by biotic community changes and deleterious effects on sensitive organisms (lichens and phytoplankton) that respond to low inputs of N (3 to 8 kilograms N per hectare per year). Streamwater nitrate concentrations are elevated in high-elevation catchments in Colorado and are unusually high in southern California and in some chaparral catchments in the southwestern Sierra Nevada. Chronic N deposition in the West is implicated in increased fire frequency in some areas and habitat alteration for threatened species. Between hotspots, N deposition is too low to cause noticeable effects or has not been studied.

Nitrogen Emissions, Deposition, and Monitoring in the Western United States

Abstract Nitrogen (N) deposition in the western United States ranges from 1 to 4 kilograms (kg) per hectare (ha) per year over much of the region to as high as 30 to 90 kg per ha per year downwind of major urban and agricultural areas. Primary N emissions sources are transportation, agriculture, and industry. Emissions of N as ammonia are about 50% as great as emissions of N as nitrogen oxides. An unknown amount of N deposition to the West Coast originates from Asia. Nitrogen deposition has increased in the West because of rapid increases in urbanization, population, distance driven, and large concentrated animal feeding operations. Studies of ecological effects suggest that emissions reductions are needed to protect sensitive ecosystem components. Deposition rates are unknown for most areas in the West, although reasonable estimates are available for sites in California, the Colorado Front Range, and central Arizona. National monitoring networks provide long-term wet deposition data and, more recently, estimated dry deposition data at remote sites. However, there is little information for many areas near emissions sources.

NITROGEN DEPOSITION IN CALIFORNIA FORESTS: A REVIEW

Atmospheric concentrations and deposition of the major nitrogenous (N) compounds and their biological effects in California forests are reviewed. Climatic characteristics of California are summarized in light of their effects on pollutant accumulation and transport. Over large areas of the state dry deposition is of greater magnitude than wet deposition due to the arid climate. However, fog deposition can also be significant in areas where seasonal fogs and N pollution sources coincide. The dominance of dry deposition is magnified in airsheds with frequent temperature inversions such as occur in the Los Angeles Air Basin. Most of the deposition in such areas occurs in summer as a result of surface deposition of nitric acid vapor (HNO3) as well as particulate nitrate (NO3-) and ammonium (NH4+). Internal uptake of gaseous N pollutants such as nitrogen dioxide (NO2), nitric oxide (NO), HNO3, peroxyacetyl nitrate (PAN), ammonia (NH3), and others provides additional N to forests. However, summer drought and subsequent lower stomatal conductance of plants tend to limit plant utilization of gaseous N. Nitrogen deposition is much greater than S deposition in California. In locations close to photochemical smog source areas, concentrations of oxidized forms of N (NO2, HNO3, PAN) dominate, while in areas near agricultural activities the importance of reduced N forms (NH3, NH4+) significantly increases. Little data from California forests are available for most of the gaseous N pollutants. Total inorganic N deposition in the most highly-exposed forests in the Los Angeles Air Basin may be as high as 25-45 kg ha(-1) year(-1). Nitrogen deposition in these highly-exposed areas has led to N saturation of chaparral and mixed conifer stands. In N saturated forests high concentrations of NO3- are found in streamwater, soil solution, and in foliage. Nitric oxide emissions from soil and foliar N:P ratios are also high in N saturated sites. Further research is needed to determine the ecological effects of chronic N deposition, and to develop appropriate management options for protecting water quality and managing plant nutrient resources in ecosystems which no longer retain excess N.

Evidence for nitrogen saturation in the San Bernardino Mountains in southern California

Abstract Elevated N deposition has occurred in the Los Angeles Basin in southern California for at least the last 40 years. Elevated streamwater NO3− fluxes and high nitric oxide (NO) fluxes from soil, indicators of N saturation, have recently been reported for chaparral watersheds exposed to chronic N deposition in the San Gabriel Mountains north/northeast of Los Angeles. A number of nutritional and edaphic parameters across a deposition gradient in the San Bernardino Mountains (SBM) support the hypothesis that the mixed conifer forest in the western end of the range is also N saturated. Concentrations of NO3− in the soil solution or in soil extracts during the summer months were 14 to 44 times higher at Camp Paivika (CP), a western high N deposition site, than at Camp Osceola (CAO) or Barton Flats (BF), eastern low-pollution sites. Accumulation of NO3− in foliage of bracken fern (Pteridium aquilinum var. pubescens Underw.) and overstory species was also much greater at CP than at CAO and a site near BF. Nitric oxide fluxes in mid-August from relatively dry soil at CP were ca. 20 times higher than for typical forests in North America. Nitrous oxide (N2O) emissions, on the other hand, were low in the SBM sites. However, emissions of NO and N2O were several-fold higher at CP than at BF, a relatively low-pollution site. High NO emissions from otherwise undisturbed and well-drained forest soils of the western US may prove useful as a diagnostic indicator of N saturation. Nitrogen mineralization was greater at CP and Dogwood (high-pollution sites) than at CAO and Heartbar (low-pollution sites). Additional indicators of N enrichment at CP compared with the low N deposition sites include: low C:N ratios in soil and foliage, high foliar N:P ratios, higher nitrification rates and high soil acidity. Lower pH and base saturation were observed in soil from two high-pollution sites compared with two low-pollution sites. In summary, high NO emissions and elevated NO3− concentrations in the soil solution and in foliage, and high foliar N:P ratios at CP, indicate N in excess of biotic demand, with potential above-normal loss of N from the ecosystem - and thus, a N-saturated condition. Model outputs from the nutrient cycling model (NuCM) agreed well with field data from the SBM on elevated soil solution NO3− concentrations, reduced soil base saturation, and lack of a growth response to increasing N deposition.

Nitrogen critical loads and management alternatives for N-impacted ecosystems in California

Empirical critical loads for N deposition effects and maps showing areas projected to be in exceedance of the critical load (CL) are given for seven major vegetation types in California. Thirty-five percent of the land area for these vegetation types (99,639 km(2)) is estimated to be in excess of the N CL. Low CL values (3-8 kg N ha(-1) yr(-1)) were determined for mixed conifer forests, chaparral and oak woodlands due to highly N-sensitive biota (lichens) and N-poor or low biomass vegetation in the case of coastal sage scrub (CSS), annual grassland, and desert scrub vegetation. At these N deposition critical loads the latter three ecosystem types are at risk of major vegetation type change because N enrichment favors invasion by exotic annual grasses. Fifty-four and forty-four percent of the area for CSS and grasslands are in exceedance of the CL for invasive grasses, while 53 and 41% of the chaparral and oak woodland areas are in exceedance of the CL for impacts on epiphytic lichen communities. Approximately 30% of the desert (based on invasive grasses and increased fire risk) and mixed conifer forest (based on lichen community changes) areas are in exceedance of the CL. These ecosystems are generally located further from emissions sources than many grasslands or CSS areas. By comparison, only 3-15% of the forested and chaparral land areas are estimated to be in exceedance of the NO(3)(-) leaching CL. The CL for incipient N saturation in mixed conifer forest catchments was 17 kg N ha(-1) yr(-1). In 10% of the CL exceedance areas for all seven vegetation types combined, the CL is exceeded by at least 10 kg N ha(-1) yr(-1), and in 27% of the exceedance areas the CL is exceeded by at least 5 kg N ha(-1) yr(-1). Management strategies for mitigating the effects of excess N are based on reducing N emissions and reducing site N capital through approaches such as biomass removal and prescribed fire or control of invasive grasses by mowing, selective herbicides, weeding or domestic animal grazing. Ultimately, decreases in N deposition are needed for long-term ecosystem protection and sustainability, and this is the only strategy that will protect epiphytic lichen communities.

Empirical and simulated critical loads for nitrogen deposition in California mixed conifer forests

Empirical critical loads (CL) for N deposition were determined from changes in epiphytic lichen communities, elevated NO(3)(-) leaching in streamwater, and reduced fine root biomass in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) at sites with varying N deposition. The CL for lichen community impacts of 3.1 kg ha(-1) year(-1) is expected to protect all components of the forest ecosystem from the adverse effects of N deposition. Much of the western Sierra Nevada is above the lichen-based CL, showing significant changes in lichen indicator groups. The empirical N deposition threshold and that simulated by the DayCent model for enhanced NO(3)(-)leaching were 17 kg N ha(-1) year(-1). DayCent estimated that elevated NO(3)(-) leaching in the San Bernardino Mountains began in the late 1950s. Critical values for litter C:N (34.1), ponderosa pine foliar N (1.1%), and N concentrations (1.0%) in the lichen Letharia vulpina ((L.) Hue) are indicative of CL exceedance.

Ozone exposure and nitrogen deposition lowers root biomass of ponderosa pine in the San Bernardino Mountains, California

Abstract Decreased root biomass in forest trees in response to anthropogenic pollutants is believed to be one of the first steps in forest health degradation. Although decreased root biomass has been observed in controlled experiments, ozone effects on mature tree roots in natural stands has not previously been documented. Here we report standing root biomass of ponderosa pine at three sites in the San Bernardino Mountains distributed along a known, long-term pollution gradient of ozone and nitrogen deposition. Trees at each site were assessed for foliar ozone injury and below-ground attributes, in addition to other environmental factors known to influence root growth. During the period of peak root growth in the spring, root biomass at the least polluted site was 6–14 times greater than that observed at the most polluted site. Known differences in climatic and edaphic factors among the sites potentially contributing to the observed response were discounted as primary contributors to the response since in most cases the site differences would have driven the patterns of root growth in the opposite direction to that observed. Differences in biotic competitive interactions, also known to affect root growth, did not explain the observed pattern for the same reason. The data suggests that elevated ozone, high nitrogen deposition, and possibly other contributing factors such as soil acidification are primarily responsible for lowering root biomass in ponderosa pine stands in the San Bernardino Mountains.

Nitrogen Deposition Effects on Mediterranean-type Ecosystems: An Ecological Assessment

We review the ecological consequences of N deposition on the five Mediterranean regions of the world. Seasonality of precipitation and fires regulate the N cycle in these water-limited ecosystems, where dry N deposition dominates. Nitrogen accumulation in soils and on plant surfaces results in peaks of availability with the first winter rains. Decoupling between N flushes and plant demand promotes losses via leaching and gas emissions. Differences in P availability may control the response to N inputs and susceptibility to exotic plant invasion. Invasive grasses accumulate as fuel during the dry season, altering fire regimes. California and the Mediterranean Basin are the most threatened by N deposition; however, there is limited evidence for N deposition impacts outside of California. Consequently, more research is needed to determine critical loads for each region and vegetation type based on the most sensitive elements, such as changes in lichen species composition and N cycling.