Hotter, longer, and more frequent global change-type drought events may profoundly impact terrestrial ecosystems by triggering widespread vegetation mortality. However, severe drought is only one component of global change, and ecological effects of drought may be compounded by other drivers, such as anthropogenic nitrogen (N) deposition and nonnative plant invasion. Elevated N deposition, for example, may reduce drought tolerance through increased plant productivity, thereby contributing to drought-induced mortality. High N availability also often favors invasive, nonnative plant species, and the loss of woody vegetation due to drought may create a window of opportunity for these invaders. We investigated the effects of multiple levels of simulated N deposition on a Mediterranean-type shrubland plant community in southern California from 2011 to 2016, a period coinciding with an extreme, multiyear drought in the region. We hypothesized that N addition would increase native shrub productivity, but that this would increase susceptibility to drought and result in increased shrub loss over time. We also predicted that N addition would favor nonnatives, especially annual grasses, leading to higher biomass and cover of these species. Consistent with these hypotheses, we found that high N availability increased native shrub canopy loss and mortality, likely due to the higher productivity and leaf area and reduced water-use efficiency we observed in shrubs subject to N addition. As native shrub cover declined, we also observed a concomitant increase in cover and biomass of nonnative annuals, particularly under high levels of experimental N deposition. Together, these results suggest that the impacts of extended drought on shrubland ecosystems may be more severe under elevated N deposition, potentially contributing to the widespread loss of native woody species and vegetation-type conversion.
In California, invasive grasses have displaced native plants, transforming much of the endemic coastal sage scrub (CSS) to nonnative grasslands. This has occurred for several reasons, including increased competitive ability of invasive grasses and long-term alterations to the soil environment, called legacy effects. Despite the magnitude of this problem, however, it is not well understood how these legacy effects have altered the soil microbial community and, indirectly, native plant restoration. We assessed the microbial composition of soils collected from an uninvaded CSS community (uninvaded soil) and a nearby 10-ha site from which the invasive grass Harding grass (Phalaris aquatica L.) was removed after 11 yr of growth (postinvasive soil). We also measured the survival rate, biomass, and length of three CSS species and P. aquatica grown in both soil types (uninvaded and postinvasive). Our findings indicate that P. aquatica may create microbial legacy effects in the soil that likely cause soil conditions inhibitory to the survival rate, biomass, and length of coastal sagebrush, but not the other two native plant species. Specifically, coastal sagebrush growth was lower in the postinvasive soil, which had more Bacteroidetes, Proteobacteria, Agrobacterium, Bradyrhizobium, Rhizobium (R. leguminosarum), Candidatus koribacter, Candidatus solibacter, and rhizophilic arbuscular mycorrhizal fungi, and fewer Planctomycetes, Acidobacteria, Nitrospira, and Rubrobacter compared with the uninvaded soil. Shifts in soil microbial community composition such as these can have important implications for restoration strategies in postinvasive sites.
Salt marshes provide storm protection to shorelines, sequester carbon (C), and mitigate coastal eutrophication. These valuable coastal ecosystems are confronted with increasing nitrogen (N) inputs from anthropogenic sources, such as agricultural runoff, wastewater, and atmospheric deposition. To inform predictions of salt marsh functioning and sustainability in the future, we characterized the response of a variety of plant, microbial, and sediment responses to a seven-level gradient of N addition in three Californian salt marshes after 7 and 14 months of N addition. The marshes showed variable responses to the experimental N gradient that can be grouped as neutral (root biomass, sediment respiration, potential carbon mineralization, and potential net nitrification), linear (increasing methane flux, decreasing potential net N mineralization, and increasing sediment inorganic N), and nonlinear (saturating aboveground plant biomass and leaf N content, and exponentially increasing sediment inorganic and organic N). The three salt marshes showed quantitative differences in most ecosystem properties and processes rates; however, the form of the response curves to N addition were generally consistent across the three marshes, indicating that the responses observed may be applicable to other marshes in the region. Only for sediment properties (inorganic and organic N pool) did the shape of the response differ significantly between marshes. Overall, the study suggests salt marshes are limited in their ability to sequester C and N with future increases in N, even without further losses in marsh area.
Abstract. Often termed "acid rain," combined nitrogen and sulfur deposition can directly and indirectly impact the condition and health of forest ecosystems. Researchers use critical loads (CLs) to describe response thresholds, and recent studies on acid-sensitive biological indicators show that forests continue to be at risk from terrestrial acidification. However, rarely are impacts translated into changes in "ecosystem services" that impact human well-being. Further, the relevance of this research to the general public is seldom communicated in terms that can motivate action to protect valuable resources. To understand how changes in biological indicators affect human well-being, we used the STEPS (Stressor-Ecological Production function-final ecosystem Services) Framework to quantitatively and qualitatively link CL exceedances to ecosystem service impacts. We specified the cause-and-effect ecological processes linking changes in biological indicators to final ecosystem services. The Final Ecosystem Goods and Services Classification System (FEGS-CS) was used within the STEPS Framework to classify the ecosystem component and the beneficiary class that uses or values the component. We analyzed two acid-sensitive tree species, balsam fir (Abies balsamea) and white ash (Fraxinus americana), that are common in northeastern USA. These well-known species provide habitat for animals and popular forest products that are relatable to a broad audience. We identified 160 chains with 10 classes of human beneficiaries for balsam fir and white ash combined, concluding that there are resources at risk that the public may value. Two stories resulting from these explorations into the cascading effects of acid rain on terrestrial resources are ideal for effective science communication: the relationship between (1) balsam fir as a popular Christmas tree and habitat for the snowshoe hare, a favorite of wildlife viewers, and (2) white ash because it is used for half of all baseball bats, fine wood products, and musical instruments. Thus, rather than focusing on biological indicators that may only be understood or appreciated by specific stakeholders or experts, this approach extends the analysis to include impacts on FEGS and humans. It also lays the foundation for developing stakeholder-specific narratives, quantitative measures of endpoints, and for conducting demand-based valuations of affected ecosystem services.
Anthropogenic nitrogen (N) deposition is known to reduce plant diversity in ecosystems worldwide; however, effects on the diversity of Mediterranean‐type ecosystems—global hotspots of biodiversity—are relatively unexplored. In California, elevated N deposition due to air pollution has a multitude of ecological effects including the facilitation of nonnative plant invasion and altered ecosystem functioning, but impacts on plant richness have been inadequately quantified. We addressed this research gap by evaluating patterns of plant richness in coastal sage scrub (CSS), a severely threatened, highly diverse Mediterranean‐type shrubland, across the Santa Monica Mountains National Recreation Area. This is the largest urban national park in the United States and experiences a strong gradient of N deposition due to its proximity to urban Los Angeles. We measured soil N, plant cover, and richness at 30 CSS sites across this gradient and used regression analyses to explore relationships between richness, N deposition, and other environmental variables. We observed significant declines in plant richness across a steep gradient of soil N availability that paralleled patterns of N deposition, primarily due to decreases in native forb species. Our analyses identified soil N as the best predictor of patterns of native forb richness, but other factors, including nonnative plant cover and aridity, may also drive reduced richness. In addition to the marked decline in the number of native forb species, increasing N deposition was also associated with lower native shrub richness per area and increased cover of nonnatives. These results highlight the threat posed by N deposition to the conservation of this already imperiled ecosystem under continued environmental change.
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