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Resilience is the ability of ecosystems to maintain function while experiencing perturbation. Globally, forests are experiencing disturbances of unprecedented quantity, type, and magnitude that may diminish resilience. Early warning signals are statistical properties of data whose increase over time may provide insights into decreasing resilience, but there have been few applications to forests. We quantified four early warning signals (standard deviation, lag-1 autocorrelation, skewness, and kurtosis) across detrended time series of multiple ecosystem state variables at the Hubbard Brook Experimental Forest, New Hampshire, USA and analyzed how these signals have changed over time. Variables were collected over periods from 25 to 55 years from both experimentally manipulated and reference areas and were aggregated to annual timesteps for analysis. Long-term (>50 year) increases in early warning signals of stream calcium, a key biogeochemical variable at the site, illustrated declining resilience after decades of acid deposition, but only in watersheds that had previously been harvested. Trends in early warning signals of stream nitrate, a critical nutrient and water pollutant, likewise exhibited symptoms of declining resilience but in all watersheds. Temporal trends in early warning signals of some of groups of trees, insects, and birds also indicated changing resilience, but this pattern differed among, and even within, groups. Overall, ∼60% of early warning signals analyzed indicated decreasing resilience. Most of these signals occurred in skewness and kurtosis, suggesting ‘flickering’ behavior that aligns with emerging evidence of the forest transitioning into an oligotrophic condition. The other ∼40% of early warning signals indicated increasing or unchanging resilience. Interpretation of early warning signals in the context of system specific knowledge is therefore essential. They can be useful indicators for some key ecosystem variables; however, uncertainties in other variables highlight the need for further development of these tools in well-studied, long-term research sites.

 

Microbial biomass is known to decrease with soil drying and to increase after rewetting due to physiological assimilation and substrate limitation under fluctuating moisture conditions, but how the effects of changing moisture conditions vary between dry and wet environments is unclear. Here, we conducted a meta‐analysis to assess the effects of elevated and reduced soil moisture on microbial biomass C (MBC) and microbial biomass N (MBN) across a broad range of forest sites between dry and wet regions. We found that the influence of both elevated and reduced soil moisture on MBC and MBN concentrations in forest soils was greater in dry than in wet regions. The influence of altered soil moisture on MBC and MBN concentrations increased significantly with the manipulation intensity but decreased with the length of experimental period, with a dramatic increase observed under a very short‐term precipitation pulse. Moisture effect did not differ between coarse‐textured and fine‐textured soils. Precipitation intensity, experimental duration, and site standardized precipitation index (dry or wet climate) were more important than edaphic factors (i.e., initial water content, bulk density, and clay content) in determining microbial biomass in response to altered moisture in forest soils. Different responses of microbial biomass in forest soils between dry and wet regions should be incorporated into models to evaluate how changes in the amount, timing, and intensity of precipitation affect soil biogeochemical processes.

 

The Hubbard Brook Experimental Forest (HBEF) was established in 1955 by the U.S. Department of Agriculture, Forest Service out of concerns about the effects of logging increasing flooding and erosion. To address this issue, within the HBEF hydrological and micrometeorological monitoring was initiated in small watersheds designated for harvesting experiments. The Hubbard Brook Ecosystem Study (HBES) originated in 1963, with the idea of using the small watershed approach to study element fluxes and cycling and the response of forest ecosystems to disturbances, such as forest management practices and air pollution. Early evidence of acid rain was documented at the HBEF and research by scientists at the site helped shape acid rain mitigation policies. New lines of investigation at the HBEF have built on the long legacy of watershed research resulting in a shift from comparing inputs and outputs and quantifying pools and fluxes to a more mechanistic understanding of ecosystem processes within watersheds. For example, hydropedological studies have shed light on linkages between hydrologic flow paths and soil development that provide valuable perspective for managing forests and understanding stream water quality. New high frequency in situ stream chemistry sensors are providing insights about extreme events and diurnal patterns that were indiscernible with traditional weekly sampling. Additionally, tools are being developed for visual and auditory data exploration and discovery by a broad audience. Given the unprecedented environmental change that is occurring, data from the small watersheds at the HBEF are more relevant now than ever and will continue to serve as a basis for sound environmental decision‐making.

 

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) 

Microbial biomass is known to decrease with soil drying and to increase after rewetting due to physiological assimilation and substrate limitation under fluctuating moisture conditions, but how the effects of moisture changes vary between dry and wet environments is unclear. Here, we conducted a meta‐analysis to assess the effects of elevated and reduced soil moisture on microbial biomass carbon (MBC) and nitrogen (MBN) across a broad range of forest sites between dry and wet regions. We found that the influence of both elevated and reduced soil moisture on MBC and MBN concentrations in forest soils was greater in dry than in wet regions. The influence of altered soil moisture on MBC and MBN concentrations increased significantly with the manipulation intensity but decreased with the length of experimental period, with a dramatic increase observed under a very short‐term precipitation pulse. Moisture effect did not differ between coarse‐ and fine‐textured soils. Precipitation intensity, experimental duration, and site standardized precipitation index (dry or wet climate) were more important than edaphic factors (i.e., initial water content, bulk density, clay content) in determining microbial biomass in response to altered moisture in forest soils. Different responses of microbial biomass in forest soils between dry and wet regions should be incorporated into models to evaluate how changes in the amount, timing and intensity of precipitation affect soil biogeochemical processes.