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Identifying which aspects of global environmental change are driving observed ecosystem process responses is a great challenge. Here, we address how long-term (10-25 year) alterations in soil moisture, and nitrogen (N) oligotrophication (i.e. decreases in soil N availability relative to plant demand), alter the production of plant-available N via net mineralization and nitrification in a northern hardwood forest. Our objectives were to determine whether soil moisture has changed over the past decade and whether N cycle processes have become less sensitive to soil moisture over time due to N oligotrophication. We used long-term data sets from several related studies to show: (i) increasing winter soil temperatures and declining summer soil moisture from late 2010 into 2024; (ii) reductions in sensitivity of N cycling rates to soil moisture, and (iii) declining moisture-adjusted N cycle processes (the ratio of rate of N process:soil moisture) over time in both summer and winter. These changes suggest continued reductions in N availability to plants in these forests, with potential effects on forest productivity and response to disturbance. 

Declining nitrogen (N) availability relative to plant demand, known as N oligotrophication, is a widespread phenomenon that has been particularly well documented in northern hardwood forests of the northeast U.S. It is hypothesized that later fall senescence contributes to this trend by increasing tree resorption of N, resulting in higher carbon:nitrogen ratios (C:N) in litterfall and reduced N availability in soil. To examine the effects of litterfall C:N on soil N cycling, we conducted a litter quality manipulation experiment comparing low C:N and high C:N litter with native litter along an elevation and aspect gradient at Hubbard Brook Experimental Forest, NH, USA. We found that potential net ammonification and mineralization rates were positively correlated with litter N and negatively correlated with litter C:N under high C:N litter, but these relationships were not present under native or low C:N litter. Differences in nitrate pools and net mineralization rates between high- and low-quality litter treatments were greater at colder sites, where native litterfall tends to have lower C:N than at low-elevation sites. Together, these results demonstrate that higher C:N litter and a warming climate may contribute to N oligotrophication through effects on microbially driven N cycling rates in organic soils. 

Snow cover is a critical factor controlling plant performance, such as survival, growth, and biomass, and vegetation cover in regions with seasonal snow (e.g., high-latitude and high elevation regions), due to its influence on the timing and length of the growing season, insulation effect during winter, and biotic and abiotic environmental factors. Therefore, changes in snow cover driven by rising temperatures and shifting precipitation patterns are expected to alter plant performance and vegetation cover. Despite the rapid increase in research on this topic in recent decades, there is still a lack of studies that quantitatively elucidate how plant performance and vegetation cover respond to shifting snow cover across snowy regions. Additionally, no comprehensive study has yet quantitatively examined these responses across regions, ecosystems, and plant functional types. Here, we conducted a meta-analysis synthesizing data from 54 snow cover manipulation studies conducted in both the field and laboratory across snowy regions to detect how plants performance and vegetation cover respond to decreased or increased snow cover. Our results demonstrate that plant survival, aboveground biomass, and belowground biomass exhibited significant decreases in response to decreased snow cover, with rates of survival having the greatest decrease. In response to increased snow cover, plant survival, growth, biomass and vegetation cover tended to increase, except for plant belowground length growth and biomass, which showed significant decreases. Additionally, our quantitative analysis of plant responses to changes in snow cover across regions, ecosystems, and plant functional types revealed that cold regions with thin snow cover, tundra and forest ecosystems, and woody species are particularly vulnerable to snow cover reduction. Overall, this study demonstrates the strong controls that snow cover exerts on plant performance, providing insights into the dynamics of snow-covered ecosystems under changing winter climatic conditions. 

Foliar resorption is a principal nutrient conservation mechanism in terrestrial vegetation that could be sensitive to ongoing changes in climate and atmospheric nitrogen (N) deposition. We quantified N resorption in northern hardwood forests along an elevation gradient of decreasing temperature and increasing soil N availability to evaluate how this critical nutrient cycling process can be expected to respond to global and regional environmental changes. Foliar N resorption proficiency (NRP) increased significantly at lower elevations for both sugar maple and American beech, the dominant species in these forests. Foliar N resorption efficiency (NRE) also decreased with increasing elevation, but only in one year. Both species exhibited strong negative relationships between NRP and soil N availability. Thus, we anticipate that with climate warming and decreasing N inputs, northern hardwood forests can be expected to exhibit stronger N conservation via foliar resorption. Both species also exhibited strong correlations between resorption efficiency of N and C, but resorption of both elements was much greater for beech than sugar maple, suggesting contrasting mechanisms of nutrient conservation between these two widespread species. 

Abstract Forest and freshwater ecosystems are tightly linked and together provide important ecosystem services, but climate change is affecting their species composition, structure, and function. Research at nine US Long Term Ecological Research sites reveals complex interactions and cascading effects of climate change, some of which feed back into the climate system. Air temperature has increased at all sites, and those in the Northeast have become wetter, whereas sites in the Northwest and Alaska have become slightly drier. These changes have altered streamflow and affected ecosystem processes, including primary production, carbon storage, water and nutrient cycling, and community dynamics. At some sites, the direct effects of climate change are the dominant driver altering ecosystems, whereas at other sites indirect effects or disturbances and stressors unrelated to climate change are more important. Long-term studies are critical for understanding the impacts of climate change on forest and freshwater ecosystems. 

BACKGROUND The availability of nitrogen (N) to plants and microbes has a major influence on the structure and function of ecosystems. Because N is an essential component of plant proteins, low N availability constrains the growth of plants and herbivores. To increase N availability, humans apply large amounts of fertilizer to agricultural systems. Losses from these systems, combined with atmospheric deposition of fossil fuel combustion products, introduce copious quantities of reactive N into ecosystems. The negative consequences of these anthropogenic N inputs—such as ecosystem eutrophication and reductions in terrestrial and aquatic biodiversity—are well documented. Yet although N availability is increasing in many locations, reactive N inputs are not evenly distributed globally. Furthermore, experiments and theory also suggest that global change factors such as elevated atmospheric CO 2 , rising temperatures, and altered precipitation and disturbance regimes can reduce the availability of N to plants and microbes in many terrestrial ecosystems. This can occur through increases in biotic demand for N or reductions in its supply to organisms. Reductions in N availability can be observed via several metrics, including lowered nitrogen concentrations ([N]) and isotope ratios (δ 15 N) in plant tissue, reduced rates of N mineralization, and reduced terrestrial N export to aquatic systems. However, a comprehensive synthesis of N availability metrics, outside of experimental settings and capable of revealing large-scale trends, has not yet been carried out. ADVANCES A growing body of observations confirms that N availability is declining in many nonagricultural ecosystems worldwide. Studies have demonstrated declining wood δ 15 N in forests across the continental US, declining foliar [N] in European forests, declining foliar [N] and δ 15 N in North American grasslands, and declining [N] in pollen from the US and southern Canada. This evidence is consistent with observed global-scale declines in foliar δ 15 N and [N] since 1980. Long-term monitoring of soil-based N availability indicators in unmanipulated systems is rare. However, forest studies in the northeast US have demonstrated decades-long decreases in soil N cycling and N exports to air and water, even in the face of elevated atmospheric N deposition. Collectively, these studies suggest a sustained decline in N availability across a range of terrestrial ecosystems, dating at least as far back as the early 20th century. Elevated atmospheric CO 2 levels are likely a main driver of declines in N availability. Terrestrial plants are now uniformly exposed to ~50% more of this essential resource than they were just 150 years ago, and experimentally exposing plants to elevated CO 2 often reduces foliar [N] as well as plant-available soil N. In addition, globally-rising temperatures may raise soil N supply in some systems but may also increase N losses and lead to lower foliar [N]. Changes in other ecosystem drivers—such as local climate patterns, N deposition rates, and disturbance regimes—individually affect smaller areas but may have important cumulative effects on global N availability. OUTLOOK Given the importance of N to ecosystem functioning, a decline in available N is likely to have far-reaching consequences. Reduced N availability likely constrains the response of plants to elevated CO 2 and the ability of ecosystems to sequester carbon. Because herbivore growth and reproduction scale with protein intake, declining foliar [N] may be contributing to widely reported declines in insect populations and may be negatively affecting the growth of grazing livestock and herbivorous wild mammals. Spatial and temporal patterns in N availability are not yet fully understood, particularly outside of Europe and North America. Developments in remote sensing, accompanied by additional historical reconstructions of N availability from tree rings, herbarium specimens, and sediments, will show how N availability trajectories vary among ecosystems. Such assessment and monitoring efforts need to be complemented by further experimental and theoretical investigations into the causes of declining N availability, its implications for global carbon sequestration, and how its effects propagate through food webs. Responses will need to involve reducing N demand via lowering atmospheric CO 2 concentrations, and/or increasing N supply. Successfully mitigating and adapting to declining N availability will require a broader understanding that this phenomenon is occurring alongside the more widely recognized issue of anthropogenic eutrophication. Intercalibration of isotopic records from leaves, tree rings, and lake sediments suggests that N availability in many terrestrial ecosystems has steadily declined since the beginning of the industrial era. Reductions in N availability may affect many aspects of ecosystem functioning, including carbon sequestration and herbivore nutrition. Shaded areas indicate 80% prediction intervals; marker size is proportional to the number of measurements in each annual mean. Isotope data: (tree ring) K. K. McLauchlan et al. , Sci. Rep. 7 , 7856 (2017); (lake sediment) G. W. Holtgrieve et al. , Science 334 , 1545–1548 (2011); (foliar) J. M. Craine et al. , Nat. Ecol. Evol. 2 , 1735–1744 (2018)