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Overstory foliage is collected in late summer from a reference forest to the west of Watershed 6 (also referred to as Bear Brook Watershed). Concentrations of C, N, P, K, Ca, Mg, and the natural abundance of N and C isotopes (delta-15N and delta-13C) in foliage are measured. These measurements, in combination with litterfall estimates of foliar biomass, allow us to estimate the pool of nutrients in foliage. They also allow us to estimate nutrient retranslocation, using measurements of leaf litterfall chemistry. Long-term measurements continue with the aim of detecting disturbances in nutrient cycling and trends in foliar chemistry over long time scales. These data were gathered as part of the Hubbard Brook Ecosystem Study (HBES). The HBES is a collaborative effort at the Hubbard Brook Experimental Forest, which is operated and maintained by the USDA Forest Service, Northern Research Station. 

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) 

The relationship between biodiversity and stability, or its inverse, temporal variability, is multidimensional and complex. Temporal variability in aggregate properties, like total biomass or abundance, is typically lower in communities with higher species diversity (i.e., the diversity–stability relationship [DSR]). At broader spatial extents, regional‐scale aggregate variability is also lower with higher regional diversity (in plant systems) and with lower spatial synchrony. However, focusing exclusively on aggregate properties of communities may overlook potentially destabilizing compositional shifts. It is not yet clear how diversity is related to different components of variability across spatial scales, nor whether regional DSRs emerge across a broad range of organisms and ecosystem types. To test these questions, we compiled a large collection of long‐term metacommunity data spanning a wide range of taxonomic groups (e.g., birds, fish, plants, invertebrates) and ecosystem types (e.g., deserts, forests, oceans). We applied a newly developed quantitative framework for jointly analyzing aggregate and compositional variability across scales. We quantified DSRs for composition and aggregate variability in local communities and metacommunities. At the local scale, more diverse communities were less variable, but this effect was stronger for aggregate than compositional properties. We found no stabilizing effect of γ‐diversity on metacommunity variability, but β‐diversity played a strong role in reducing compositional spatial synchrony, which reduced regional variability. Spatial synchrony differed among taxa, suggesting differences in stabilization by spatial processes. However, metacommunity variability was more strongly driven by local variability than by spatial synchrony. Across a broader range of taxa, our results suggest that high γ‐diversity does not consistently stabilize aggregate properties at regional scales without sufficient spatial β‐diversity to reduce spatial synchrony.

 

Synchrony is broadly important to population and community dynamics due to its ubiquity and implications for extinction dynamics, system stability, and species diversity. Investigations of synchrony in community ecology have tended to focus on covariance in the abundances of multiple species in a single location. Yet, the importance of regional environmental variation and spatial processes in community dynamics suggests that community properties, such as species richness, could fluctuate synchronously across patches in a metacommunity, in an analog of population spatial synchrony. Here, we test the prevalence of this phenomenon and the conditions under which it may occur using theoretical simulations and empirical data from 20 marine and terrestrial metacommunities. Additionally, given the importance of biodiversity for stability of ecosystem function, we posit that spatial synchrony in species richness is strongly related to stability. Our findings show that metacommunities often exhibit spatial synchrony in species richness. We also found that richness synchrony can be driven by environmental stochasticity and dispersal, two mechanisms of population spatial synchrony. Richness synchrony also depended on community structure, including species evenness and beta diversity. Strikingly, ecosystem stability was more strongly related to richness synchrony than to species richness itself, likely because richness synchrony integrates information about community processes and environmental forcing. Our study highlights a new approach for studying spatiotemporal community dynamics and emphasizes the spatial dimensions of community dynamics and stability.