Functional trait-based restoration alters nutrient cycling and invasion rates in Hawaiian lowland wet forest.

Many degraded ecosystems have altered nutrient dynamics due to invaders' possessing a suite of traits that allow them to both outcompete native species and alter the environment. In ecosystems where invasive species have increased nutrient turnover rates, it can be difficult to reduce nutrient availability. This study examined whether a functional trait-based restoration approach involving the planting of species with conservative nutrient-use traits could slow rates of nutrient cycling and consequently reduce rates of invasion. We examined a functional trait restoration initiative in a heavily invaded lowland wet forest site in Hilo, Hawai'i. Native and introduced species were chosen to create four experimental hybrid forest communities, in comparison to the invaded forest, with a factorial design in which communities varied in rates of carbon turnover (slow or moderate [SLOW, MOD]), and in the relationship of species in trait space (redundant or complementary [RED, COMP]). After the first 5 years, we evaluated community-level outcomes related to nutrient cycling: carbon (C), nitrogen (N), and phosphorus (P) via litterfall, litter decomposition, and outplant productivity and rates of invasion. We found that (1) regardless of treatment, the experimental communities had low rates of nutrient cycling through litterfall relative to the invaded reference forest, (2) the MOD communities had greater nutrient release via litterfall than the SLOW communities, (3) introduced species had greater nutrient release than native species in the two MOD experimental communities, and (4) within treatments, there was a positive relationship between nutrient levels and outplant basal area, but outplant basal area was negatively associated with rates of invasion. The negative relationships among basal area and weed invasion, particularly for the two COMP treatments, suggest species existing in different parts of trait space may help confer some degree of invasion resistance. The diversification of trait space was facilitated by the use of introduced species, a new concept in Hawaiian forest management. Although challenges remain in endeavors to restore this heavily degraded ecosystem, this study provides evidence that functional trait-based restoration approaches using carefully crafted hybrid communities can reduce rates of nutrient cycling and invasion in order to reach management goals.

Modeling global oceanic nitrogen deposition from food systems and its mitigation potential by reducing overuse of fertilizers.

Growing population and consumption pose unprecedented demands on food production. However, ammonia emissions mainly from food systems increase oceanic nitrogen deposition contributing to eutrophication. Here, we developed a long-term oceanic nitrogen deposition dataset (1970 to 2018) with updated ammonia emissions from food systems, evaluated the impact of ammonia emissions on oceanic nitrogen deposition patterns, and discussed the potential impact of nitrogen fertilizer overuse. Based on the chemical transport modeling approach, oceanic ammonia-related nitrogen deposition increased by 89% globally between 1970 and 2018, and now, it exceeds oxidized nitrogen deposition by over 20% in coastal regions including China Sea, India Coastal, and Northeastern Atlantic Shelves. Approximately 38% of agricultural nitrogen fertilizer was excessive, which corresponds to 15% of global oceanic ammonia-related nitrogen deposition. Policymakers and water quality managers need to pay increasingly more attention to ammonia associated with food production if the goal of reducing coastal nitrogen pollution is to be achieved for Sustainable Development Goals.

Manganese Oxidation States in Volcanic Soils across Annual Rainfall Gradients.

Manganese (Mn) exists as Mn(II), Mn(III), or Mn(IV) in soils, and the Mn oxidation state controls the roles of Mn in numerous environmental processes. However, the variations of Mn oxidation states with climate remain unknown. We determined the Mn oxidation states in highly weathered bulk volcanic soils (primary minerals free) across two rainfall gradients covering mean annual precipitation (MAP) of 0.25-5 m in the Hawaiian Islands. With increasing MAP, the soil redox conditions generally shifted from oxic to suboxic and to anoxic despite fluctuating at each site; concurrently, the proportions of Mn(IV) and Mn(II) decreased and increased, respectively. Mn(III) was low at both low and high MAP, but accumulated substantially, up to 80% of total Mn, in soils with prevalent suboxic conditions at intermediate MAP. Mn(III) was likely hosted in Mn(III,IV) and iron(III) oxides or complexed with organic matter, and its distribution among these hosts varied with soil redox potentials and soil pH. Soil redox conditions and rainfall-driven leaching jointly controlled exchangeable Mn(II) in soils, with its concentration peaking at intermediate MAP. The Mn redox chemistry was at disequilibrium, with the oxidation states correlating with long-term average soil redox potentials better than with soil pH. The soil redox conditions likely fluctuated between oxic and anoxic conditions more frequently at intermediate than at low and high MAP, creating biogeochemical hot spots where Mn, Fe, and other redox-sensitive elements may be actively cycled.

Water balance affects foliar and soil nutrients differently.

Water balance influences soil development, and consequently plant communities, by driving weathering of soil minerals and leaching of plant nutrients from the soil. Along gradients in water balance, soils exhibit process domains where chemical properties are relatively stable punctuated by pedogenic thresholds where soil chemical properties change rapidly with little additional change in water balance. We ask if plant macronutrient concentrations in leaves also exhibit non-linear trends along water balance gradients, and if so, how these non-linearities relate to those in soils. We analyze foliar nutrient concentrations and foliar N:P ratios from eight species that span a range of growth forms along three water balance gradients (three of the species are found on multiple gradients). The gradients are located on basaltic substrate of different ages and have previously been characterized by studies on soil development. We find that maximum concentrations of foliar macronutrients occur at an intermediate water balance. As with soil nutrients, time mediates the effect of water balance on foliar nutrients, such that plants on older soils attain maximum nutrient concentrations at a lower water balance. On both a young, 20 ky and an old, 4100 ky water balance gradient, foliar nutrients reach peak concentrations at a water balance greater than the threshold for depletion of rock-derived nutrients in surface soils. Our findings suggest that plant acquisition of essential nutrients is imperfectly predicted by overall soil nutrient availability because the regulation of internal nutrient pools by plants makes nutrient pools within leaves partially independent of soil nutrient availability.

Exploring global changes in agricultural ammonia emissions and their contribution to nitrogen deposition since 1980.

SignificanceAgricultural systems are already major forces of ammonia pollution and environmental degradation. How agricultural ammonia emissions affect the spatio-temporal patterns of nitrogen deposition and where to target future mitigation efforts, remains poorly understood. We develop a substantially complete and coherent agricultural ammonia emissions dataset in nearly recent four decades, and evaluate the relative role of reduced nitrogen in total nitrogen deposition in a spatially explicit way. Global reduced nitrogen deposition has grown rapidly, and will occupy a greater dominant position in total nitrogen deposition without future ammonia regulations. Recognition of agricultural ammonia emissions on nitrogen deposition is critical to formulate effective policies to address ammonia related environmental challenges and protect ecosystems from excessive nitrogen inputs.

Foliar ẟ15N patterns in legumes and non-N fixers across a climate gradient, Hawai'i Island, USA.

Recent studies from the Hawaiian Islands showed that pedogenic thresholds demarcate domains in which rock-derived nutrient dynamics remain similar across wide variations in rainfall. These thresholds appear related to certain aspects of N cycling, but the degree to which they correspond to patterns of biological N fixation (BNF)-the dominant input of N into less-managed ecosystems-remains unclear. We measured aboveground plant biomass, foliar nutrient concentrations, and foliar δ15N along a climate gradient on ~ 150,000-year-old basaltic substrate to characterize foliar N sources and spatially relate them to soil nutrients. Patterns in legume δ15N correspond to known pedogenic thresholds along the rainfall gradient, with low δ15N values (~ 0 to - 2‰) occurring in the dry, biologically inactive domain and the wet, highly weathered domain. Elevated δ15N in the middle, fertile domain suggests a greater reliance of legumes on soil N where it has accumulated over time. Non-legume face N deficiencies throughout most of the gradient while legumes maintain low C:N ratios via symbiotic BNF. However, legume abundance declines outside the fertile domain, limiting ecosystem N inputs. Breakpoints in legume δ15N data suggest that P (and potentially other nutrients) limits BNF and, by extension, legume abundance in wet region. Nutrients may also constrain legume abundance in the dry domain, but pedogenic effects could not be isolated from climatic constraints at the dry sites. We conclude that pedogenic thresholds defined by climate can be informative of foliar δ15N patterns in cases where legumes are not directly constrained by climate, land use, or other external factors.

Soil-Food-Environment-Health Nexus for Sustainable Development.

Changes in soil properties and processes can influence food and environmental quality, thus, affecting human health and welfare through biogeochemical cascades among soil, food, environment, and human health. However, because many soil properties change much more slowly than do management practices and pollution to soil, the legacy of past influences on soil can have long-term effects on both human health and sustainability. It is essential and urgent to manage soils for health and sustainability through building the soil-food-environment-health nexus.

Nitrogen deposition accelerates soil carbon sequestration in tropical forests.

Terrestrial ecosystem carbon (C) sequestration plays an important role in ameliorating global climate change. While tropical forests exert a disproportionately large influence on global C cycling, there remains an open question on changes in below-ground soil C stocks with global increases in nitrogen (N) deposition, because N supply often does not constrain the growth of tropical forests. We quantified soil C sequestration through more than a decade of continuous N addition experiment in an N-rich primary tropical forest. Results showed that long-term N additions increased soil C stocks by 7 to 21%, mainly arising from decreased C output fluxes and physical protection mechanisms without changes in the chemical composition of organic matter. A meta-analysis further verified that soil C sequestration induced by excess N inputs is a general phenomenon in tropical forests. Notably, soil N sequestration can keep pace with soil C, based on consistent C/N ratios under N additions. These findings provide empirical evidence that below-ground C sequestration can be stimulated in mature tropical forests under excess N deposition, which has important implications for predicting future terrestrial sinks for both elevated anthropogenic CO2 and N deposition. We further developed a conceptual model hypothesis depicting how soil C sequestration happens under chronic N deposition in N-limited and N-rich ecosystems, suggesting a direction to incorporate N deposition and N cycling into terrestrial C cycle models to improve the predictability on C sink strength as enhanced N deposition spreads from temperate into tropical systems.

Diversity of putative ericoid mycorrhizal fungi increases with soil age and progressive phosphorus limitation across a 4.1-million-year chronosequence.

Ericaceous plants rely on ericoid mycorrhizal fungi for nutrient acquisition. However, the factors that affect the composition and structure of fungal communities associated with the roots of ericaceous plants remain largely unknown. Here, we use a 4.1-million-year (myr) soil chronosequence in Hawaii to test the hypothesis that changes in nutrient availability with soil age determine the diversity and species composition of fungi associated with ericoid roots. We sampled roots of a native Hawaiian plant, Vaccinium calycinum, and used DNA metabarcoding to quantify changes in fungal diversity and community composition. We also used a fertilization experiment at the youngest and oldest sites to assess the importance of nutrient limitation. We found an increase in diversity and a clear pattern of species turnover across the chronosequence, driven largely by putative ericoid mycorrhizal fungi. Fertilization with nitrogen at the youngest site and phosphorus at the oldest site reduced fungal diversity, suggesting a direct role of nutrient limitation. Our results also reveal the presence of novel fungal species associated with Hawaiian Ericaceae and suggest a greater importance of phosphorus availability for communities of ericoid mycorrhizal fungi than is generally assumed.

Changes in belowground biodiversity during ecosystem development.

Belowground organisms play critical roles in maintaining multiple ecosystem processes, including plant productivity, decomposition, and nutrient cycling. Despite their importance, however, we have a limited understanding of how and why belowground biodiversity (bacteria, fungi, protists, and invertebrates) may change as soils develop over centuries to millennia (pedogenesis). Moreover, it is unclear whether belowground biodiversity changes during pedogenesis are similar to the patterns observed for aboveground plant diversity. Here we evaluated the roles of resource availability, nutrient stoichiometry, and soil abiotic factors in driving belowground biodiversity across 16 soil chronosequences (from centuries to millennia) spanning a wide range of globally distributed ecosystem types. Changes in belowground biodiversity during pedogenesis followed two main patterns. In lower-productivity ecosystems (i.e., drier and colder), increases in belowground biodiversity tracked increases in plant cover. In more productive ecosystems (i.e., wetter and warmer), increased acidification during pedogenesis was associated with declines in belowground biodiversity. Changes in the diversity of bacteria, fungi, protists, and invertebrates with pedogenesis were strongly and positively correlated worldwide, highlighting that belowground biodiversity shares similar ecological drivers as soils and ecosystems develop. In general, temporal changes in aboveground plant diversity and belowground biodiversity were not correlated, challenging the common perception that belowground biodiversity should follow similar patterns to those of plant diversity during ecosystem development. Taken together, our findings provide evidence that ecological patterns in belowground biodiversity are predictable across major globally distributed ecosystem types and suggest that shifts in plant cover and soil acidification during ecosystem development are associated with changes in belowground biodiversity over centuries to millennia.

Nitrogen dynamics along a climate gradient on geologically old substrate, Kaua'i, Hawai'i.

We evaluated N dynamics on a climate gradient on old (> 4 million year) basaltic substrate on the Island of Kaua'i, Hawai'i, to evaluate the utility of pedogenic thresholds and soil process domains for understanding N cycling in terrestrial ecosystems. Studies of nitrogen dynamics on the climate gradient on a younger basaltic substrate (~ 150,000 year) had found a good match between soil process domains and N cycling processes. Here we measured net N mineralization and nitrification by incubation, and δ15N of total soil N, to determine whether the soil process domains on the older gradient were equally useful for interpreting N cycling and thereby to explore the general utility of the approach. Net N mineralization varied from 0 to 1.7 mg kg-1 d-1 across the old Kaua'i gradient, and δ15N varied from + 3 to + 11 ο/οο, both ranges similar to those on the younger substrate. However, while the pattern of variation with climate was similar for δ15N, the highest rates of mineralization on the old gradient occurred where forests were dominated by the native N fixer Acacia koa. This occurred in sites wetter than the process domain associated with high net N mineralization on the gradient on younger substrate. We conclude that soil process domains based on rock-derived nutrients are not always useful for evaluating N dynamics, especially where the distribution of biological N fixers is controlled by factors other than rock-derived nutrients.

Quantitative Analysis of Pedogenic Thresholds and Domains in Volcanic Soils.

Pedogenic thresholds describe where soil properties or processes change in an abrupt/nonlinear fashion in response to small changes in environmental forcing. Contrastingly, soil process domains refer to the space between thresholds where soil properties are either unchanged, or change gradually, across a broad range of environmental forcing. Here, we test quantitatively for the presence of thresholds in patterns of soil properties across a climatic gradient on soils developed from ~20 ky old basaltic substrate on the Island of Hawai'i. From multiple soil properties, we quantitatively identified a threshold at ~750 mm/y of water balance (precipitation minus potential evapotranspiration), delineating the upper water balance boundary of soil fertility in these soils. From the threshold in the ratio of exchangeable Ca to total Ca we identified the lower water balance boundary of soil fertility in these soils at -1000 mm/y, however this threshold was qualitatively described as it lies near the limit of the climate gradient data where the statistical approach can not be applied. These two results represent the first time that pedogenic thresholds have been identified using statistically rigorous methods and the limitations of said methods, respectively. Comparing the 20 ky soils to soils that developed on basaltic substrates of 1.2 ky, 7.5 ky, 150 ky, and 4100 ky in a time-climate matrix, we found that our quantitative analysis supports previous qualitatively identified thresholds in the soils developed from older substrates. We also identified the 20 ky as the transition from kinetic to supply limitation for plant nutrients in soil in this system.

Combining spectroscopic and isotopic techniques gives a dynamic view of phosphorus cycling in soil.

Current understanding of phosphorus (P) cycling in soils can be enhanced by integrating previously discrete findings concerning P speciation, exchange kinetics, and the underlying biological and geochemical processes. Here, we combine sequential extraction with P K-edge X-ray absorption spectroscopy and isotopic methods (33P and 18O in phosphate) to characterize P cycling on a climatic gradient in Hawaii. We link P pools to P species and estimate the turnover times for commonly considered P pools. Dissolved P turned over in seconds, resin-extractable P in minutes, NaOH-extractable inorganic P in weeks to months, and HCl-extractable P in years to millennia. Furthermore, we show that in arid-zone soils, some primary mineral P remains even after 150 ky of soil development, whereas in humid-zone soils of the same age, all P in all pools has been biologically cycled. The integrative information we provide makes possible a more dynamic, process-oriented conceptual model of P cycling in soils.

Policy distortions, farm size, and the overuse of agricultural chemicals in China.

Understanding the reasons for overuse of agricultural chemicals is critical to the sustainable development of Chinese agriculture. Using a nationally representative rural household survey from China, we found that farm size is a strong factor that affects the use intensity of agricultural chemicals across farms in China. Statistically, a 1% increase in farm size is associated with a 0.3% and 0.5% decrease in fertilizer and pesticide use per hectare (P < 0.001), respectively, and an almost 1% increase in agricultural labor productivity, while it only leads to a statistically insignificant 0.02% decrease in crop yields. The same pattern was also found using other independently collected data sources from China and an international panel analysis of 74 countries from the 1960s to the 2000s. While economic growth has been associated with increasing farm size in many other countries, in China this relationship has been distorted by land and migration policies, leading to the persistence of small farm size in China. Removing these distortions would decrease agricultural chemical use by 30-50% and the environmental impact of those chemicals by 50% while doubling the total income of all farmers including those who move to urban areas. Removing policy distortions is also likely to complement other remedies to the overuse problem, such as easing farmer's access to modern technologies and knowledge, and improving environmental regulation and enforcement.

Plant acclimation to long-term high nitrogen deposition in an N-rich tropical forest.

Anthropogenic nitrogen (N) deposition has accelerated terrestrial N cycling at regional and global scales, causing nutrient imbalance in many natural and seminatural ecosystems. How added N affects ecosystems where N is already abundant, and how plants acclimate to chronic N deposition in such circumstances, remains poorly understood. Here, we conducted an experiment employing a decade of N additions to examine ecosystem responses and plant acclimation to added N in an N-rich tropical forest. We found that N additions accelerated soil acidification and reduced biologically available cations (especially Ca and Mg) in soils, but plants maintained foliar nutrient supply at least in part by increasing transpiration while decreasing soil water leaching below the rooting zone. We suggest a hypothesis that cation-deficient plants can adjust to elevated N deposition by increasing transpiration and thereby maintaining nutrient balance. This result suggests that long-term elevated N deposition can alter hydrological cycling in N-rich forest ecosystems.

Convergence and contrast in the community structure of Bacteria, Fungi and Archaea along a tropical elevation-climate gradient.

Changes in species richness along climatological gradients have been instrumental in developing theories about the general drivers of biodiversity. Previous studies on microbial communities along climate gradients on mountainsides have revealed positive, negative and neutral richness trends. We examined changes in richness and composition of Fungi, Bacteria and Archaea in soil along a 50-1000 m elevation, 280-3280 mm/yr precipitation gradient in Hawai'i. Soil properties and their drivers are exceptionally well understood along this gradient. All three microbial groups responded strongly to the gradient, with community ordinations being similar along axes of environmental conditions (pH, rainfall) and resource availability (nitrogen, phosphorus). However, the form of the richness-climate relationship varied between Fungi (positive linear), Bacteria (unimodal) and Archaea (negative linear). These differences were related to resource-ecology and limiting conditions for each group, with fungal richness increasing most strongly with soil carbon, ammonia-oxidizing Archaea increasing with nitrogen mineralization rate, and Bacteria increasing with both carbon and pH. Reponses to the gradient became increasingly variable at finer taxonomic scales and within any taxonomic group most individual OTUs occurred in narrow climate-elevation ranges. These results show that microbial responses to climate gradients are heterogeneous due to complexity of underlying environmental changes and the diverse ecologies of microbial taxa.

Controls of nitrogen cycling evaluated along a well-characterized climate gradient.

The supply of nitrogen (N) constrains primary productivity in many ecosystems, raising the question "what controls the availability and cycling of N"? As a step toward answering this question, we evaluated N cycling processes and aspects of their regulation on a climate gradient on Kohala Volcano, Hawaii, USA. The gradient extends from sites receiving <300 mm/yr of rain to those receiving >3,000 mm/yr, and the pedology and dynamics of rock-derived nutrients in soils on the gradient are well understood. In particular, there is a soil process domain at intermediate rainfall within which ongoing weathering and biological uplift have enriched total and available pools of rock-derived nutrients substantially; sites at higher rainfall than this domain are acid and infertile as a consequence of depletion of rock-derived nutrients, while sites at lower rainfall are unproductive and subject to wind erosion. We found elevated rates of potential net N mineralization in the domain where rock-derived nutrients are enriched. Higher-rainfall sites have low rates of potential net N mineralization and high rates of microbial N immobilization, despite relatively high rates of gross N mineralization. Lower-rainfall sites have moderately low potential net N mineralization, relatively low rates of gross N mineralization, and rates of microbial N immobilization sufficient to sequester almost all the mineral N produced. Bulk soil δ15 N also varied along the gradient, from +4‰ at high rainfall sites to +14‰ at low rainfall sites, indicating differences in the sources and dynamics of soil N. Our analysis shows that there is a strong association between N cycling and soil process domains that are defined using soil characteristics independent of N along this gradient, and that short-term controls of N cycling can be understood in terms of the supply of and demand for N.

Integrated reactive nitrogen budgets and future trends in China.

Reactive nitrogen (Nr) plays a central role in food production, and at the same time it can be an important pollutant with substantial effects on air and water quality, biological diversity, and human health. China now creates far more Nr than any other country. We developed a budget for Nr in China in 1980 and 2010, in which we evaluated the natural and anthropogenic creation of Nr, losses of Nr, and transfers among 14 subsystems within China. Our analyses demonstrated that a tripling of anthropogenic Nr creation was associated with an even more rapid increase in Nr fluxes to the atmosphere and hydrosphere, contributing to intense and increasing threats to human health, the sustainability of croplands, and the environment of China and its environs. Under a business as usual scenario, anthropogenic Nr creation in 2050 would more than double compared with 2010 levels, whereas a scenario that combined reasonable changes in diet, N use efficiency, and N recycling could reduce N losses and anthropogenic Nr creation in 2050 to 52% and 64% of 2010 levels, respectively. Achieving reductions in Nr creation (while simultaneously increasing food production and offsetting imports of animal feed) will require much more in addition to good science, but it is useful to know that there are pathways by which both food security and health/environmental protection could be enhanced simultaneously.