Biogeochemistry research needs: observations from the ecosystem studies program of The National Science Foundation

This comment presents current and emerging areas of research which the U.S. National Science Foundation believes have special promise for advancing ecosystem science. These areas are: (1) major element cycle interactions; (2) trace element and organic compound controls on ecosystem dynamics; (3) the role of consumers on ecosystem dynamics; (4) ecosystem dynamics in contrasting environments; (5) landscape ecology; and (6) enhancement of theory and methodology. The Ecosystem Studies Program of the National Science Foundation has identified these as areas to be emphasized in its long range planning program (revised March, 1985). Biogeochemical studies are emphasized in many of these research priority areas.     

Biogeochemistry of Late Paleozoic North American brachiopods and secular variation of seawater composition

Sr/Ca ratios in modern brachiopod shells reflect variations in ambient seawater, whereas their Na contents show no relationship with water depth or habitat. Their Mn and Fe contents are controlled, in part, by leaching of these elements from oxide coatings or the low input/sedimentation rate of detrital material into depositional areas such as Quatsino Sound. For most Carboniferous brachiopods from North America, the Mn and Fe contents are similar to those recorded by their Recent counterparts. The high Mn and Fe contents in the brachiopods from shales suggest several possibilities for these levels. One possibility is the leaching of Mn and Fe from oxide coatings/matrix which was not completely removed in the cleaning process, or the high levels in part reflect unusual depositional conditions (some degree of anoxia) for the local shaly environments. The Sr/Ca ratio of brachiopods and, by inference, complementary seawater, did not vary significantly during the Carboniferous. The Sr/Ca minimum observed in brachiopods of Mississippian age coincides with a dip in the ⁸⁷Sr/⁸⁶Sr curve and correlates with the Hercynian orogeny. This is attributed to the cycling of seawater through mid-ocean ridge basalts, and postulated exchange reactions account for variation in the composition of seawater-Ca. The unidirectional trend of heavier δ¹³C values from the Devonian to the Permian is intricately coupled with the evolution of the terrestrial biomass. In addition to expansion of terrestrial plants, burial of reduced carbon in the form of coal (organic matter) contributed to the observed shift. The start of the Permo-Pennsylvanian glaciation is marked by a negative excursion of the secular carbon trend, which is linked to weathering of reduced carbon and its return to the oceanic reservoir with its oxidized carbon. The oxygen isotope values reflect the unidirectional trend towards higher values of the carbon data with decreasing geologic age. Negative excursions of the trend may be related to extensive weathering of terrestrial and submarine rocks, whereas positive excursions may be related to hydrothermal alteration of submarine rocks and dehydration of oceanic crust during times of active sea-floor spreading. Oxygen-calculated water temperatures of unaltered brachiopod material are unrealistically high for all of the Devonian, and the Chesterian-Meramecian, Desmoinesian-Missourian, and Artinskian Epochs. During these times maximum water temperatures of 42° to 56°C are well above the thermal threshold of protein denaturation. This process, which is lethal to most higher organisms, demands an adjustment in oxygen of -2.5%. for samples older than Missourian, and of -1.250%. for samples spanning the Missourian-Artinskian interval. With these adjustments and salinity considerations made prior to calculations, water temperatures become reasonable for the Late Paleozoic epeiric, tropical seas of North America.     

Resource quality affects carbon cycling in deep-sea sediments

Deep-sea sediments cover ∼70% of Earth''s surface and represent the largest interface between the biological and geological cycles of carbon. Diatoms and zooplankton faecal pellets naturally transport organic material from the upper ocean down to the deep seabed, but how these qualitatively different substrates affect the fate of carbon in this permanently cold environment remains unknown. We added equal quantities of ¹³C-labelled diatoms and faecal pellets to a cold water (−0.7 °C) sediment community retrieved from 1080 m in the Faroe-Shetland Channel, Northeast Atlantic, and quantified carbon mineralization and uptake by the resident bacteria and macrofauna over a 6-day period. High-quality, diatom-derived carbon was mineralized >300% faster than that from low-quality faecal pellets, demonstrating that qualitative differences in organic matter drive major changes in the residence time of carbon at the deep seabed. Benthic bacteria dominated biological carbon processing in our experiments, yet showed no evidence of resource quality-limited growth; they displayed lower growth efficiencies when respiring diatoms. These effects were consistent in contrasting months. We contend that respiration and growth in the resident sediment microbial communities were substrate and temperature limited, respectively. Our study has important implications for how future changes in the biochemical makeup of exported organic matter will affect the balance between mineralization and sequestration of organic carbon in the largest ecosystem on Earth.     

The forest–alpine ecotone: a multi-scale approach to spatial and temporal dynamics of treeline change at Niwot Ridge

Background: Current understanding of treeline or forest-alpine ecotone (FAE) dynamics does not fully explain past and present FAE patterns and their underlying processes, nor allow prediction of their response to climate change.

Aims: We address the overarching hypothesis that the FAE is a mosaic of distinct landscape units of vegetation and landforms that result in differential responses to climate change. We focus on climate-related, landscape and vegetation characteristics, but also consider the effect of landscape heterogeneity on biogeochemistry and overall resilience of the FAE to climate change.

Results: There are three distinct FAE land units at Niwot Ridge, generated by different interactions of climate with vegetation, landforms and topography. Within these FAEs, a process of self-organisation takes place from organism to patch to landscape scales, and is modulated by positive and negative feedback loops along an elevation gradient. The underlying controls cannot be attributed solely to temperature, but to a combination of interactions along a physical/biotic gradient.

Conclusions: FAE dynamics result from interactions among mechanisms and processes at the microsite, patch and landscape scales: (1) tree persistence; (2) forest patch establishment; (3) drivers of patch forest configurations and (4) resilience, increasing along a gradient of biotic control.     

Proton production from nitrogen transformation drives stream export of base cations in acid-sensitive forested watersheds

The biogeochemical cycles of nitrogen (N) and base cations (BCs), (i.e., K⁺, Na⁺, Ca²⁺, and Mg²⁺), play critical roles in plant nutrition and ecosystem function. Empirical correlations between large experimental N fertilizer additions to forest ecosystems and increased BCs loss in stream water are well demonstrated, but the mechanisms driving this coupling remain poorly understood. We hypothesized that protons generated through N transformation (PPR_N)—quantified as the balance of NH₄⁺ (H⁺ source) and NO₃⁻ (H⁺ sink) in precipitation versus the stream output will impact BCs loss in acid-sensitive ecosystems. To test this hypothesis, we monitored precipitation input and stream export of inorganic N and BCs for three years in an acid-sensitive forested watershed in a granite area of subtropical China. We found the precipitation input of inorganic N (17.71 kg N ha⁻¹ year⁻¹ with 54% as NH₄⁺–N) was considerably higher than stream exported inorganic N (5.99 kg N ha⁻¹ year⁻¹ with 83% as NO₃⁻–N), making the watershed a net N sink. The stream export of BCs (151, 1518, 851, and 252 mol ha⁻¹ year⁻¹ for K⁺, Na⁺, Ca²⁺, and Mg²⁺, respectively) was positively correlated (r = 0.80, 0.90, 0.84, and 0.84 for K⁺, Na⁺, Ca²⁺, and Mg²⁺ on a monthly scale, respectively, P < 0.001, n = 36) with PPR_N (389 mol ha⁻¹ year⁻¹) over the three years, suggesting that PPR_N drives loss of BCs in the acid-sensitive ecosystem. A global meta-analysis of 15 watershed studies from non-calcareous ecosystems further supports this hypothesis by showing a similarly strong correlation between ∑BCs output and PPR_N (r = 0.89, P < 0.001, n = 15), in spite of the pronounced differences in environmental settings. Collectively, our results suggest that N transformations rather than anions (NO₃⁻ and/or SO₄²⁻) leaching specifically, are an important mediator of BCs loss in acid-senstive ecosystems. Our study provides the first definitive evidence that the chronic N deposition and subsequent transformation within the watershed drive stream export of BCs through proton production in acid-sensitive ecosystems, irrespective of their current relatively high N retention. Our findings suggest the N-transformation-based proton production can be used as an indicator of watershed outflow quality in the acid-sensitive ecosystems.     

Altered dynamics of forest recovery under a changing climate

Forest regeneration following disturbance is a key ecological process, influencing forest structure and function, species assemblages, and ecosystem–climate interactions. Climate change may alter forest recovery dynamics or even prevent recovery, triggering feedbacks to the climate system, altering regional biodiversity, and affecting the ecosystem services provided by forests. Multiple lines of evidence – including global‐scale patterns in forest recovery dynamics; forest responses to experimental manipulation of CO₂, temperature, and precipitation; forest responses to the climate change that has already occurred; ecological theory; and ecosystem and earth system models – all indicate that the dynamics of forest recovery are sensitive to climate. However, synthetic understanding of how atmospheric CO₂ and climate shape trajectories of forest recovery is lacking. Here, we review these separate lines of evidence, which together demonstrate that the dynamics of forest recovery are being impacted by increasing atmospheric CO₂ and changing climate. Rates of forest recovery generally increase with CO₂, temperature, and water availability. Drought reduces growth and live biomass in forests of all ages, having a particularly strong effect on seedling recruitment and survival. Responses of individual trees and whole‐forest ecosystems to CO₂ and climate manipulations often vary by age, implying that forests of different ages will respond differently to climate change. Furthermore, species within a community typically exhibit differential responses to CO₂ and climate, and altered community dynamics can have important consequences for ecosystem function. Age‐ and species‐dependent responses provide a mechanism by which climate change may push some forests past critical thresholds such that they fail to recover to their previous state following disturbance. Altered dynamics of forest recovery will result in positive and negative feedbacks to climate change. Future research on this topic and corresponding improvements to earth system models will be a key to understanding the future of forests and their feedbacks to the climate system.     

Diagnosing and assessing uncertainties of terrestrial ecosystem models in a multimodel ensemble experiment: 2. Carbon balance

This paper examines carbon stocks and their relative balance in terrestrial ecosystems simulated by Biome‐BGC, LPJ, and CASA in an ensemble model experiment conducted using the Terrestrial Observation and Prediction System. We developed the Hierarchical Framework for Diagnosing Ecosystem Models to separate the simulated biogeochemistry into a cascade of functional tiers and examine their characteristics sequentially. The analyses indicate that the simulated biomass is usually two to three times higher in Biome‐BGC than LPJ or CASA. Such a discrepancy is mainly induced by differences in model parameters and algorithms that regulate the rates of biomass turnover. The mean residence time of biomass in Biome‐BGC is estimated to be 40–80 years in temperate/moist climate regions, while it mostly varies between 5 and 30 years in CASA and LPJ. A large range of values is also found in the simulated soil carbon. The mean residence time of soil carbon in Biome‐BGC and LPJ is ～200 years in cold regions, which decreases rapidly with increases of temperature at a rate of ～10 yr °C^?1. Because long‐term soil carbon pool is not simulated in CASA, its corresponding mean residence time is only about 10–20 years and less sensitive to temperature. Another key factor that influences the carbon balance of the simulated ecosystem is disturbance caused by wildfire, for which the algorithms vary among the models. Because fire emissions are balanced by net ecosystem production (NEP) at steady states, magnitudes, and spatial patterns of NEP vary significantly as well. Slight carbon imbalance may be left by the spin‐up algorithm of the models, which adds uncertainty to the estimated carbon sources or sinks. Although these results are only drawn on the tested model versions, the developed methodology has potential for other model exercises.     

Insights into the spatiotemporal component of feeding ecology: an isotopic approach for conservation management sciences

Aim Bringing new approaches to trace spatiotemporal variations in animals’ feeding ecology, which is fundamental for wildlife management and conservation since the accessibility of animals to trophic resources plays a key role in the dynamics of populations and metapopulations. Location Western Mediterranean coast. Methods The analysis of naturally occurring stable isotopes constitutes an exceptional approach to assess variations in the trophic ecology of species within the spatiotemporal dimensions. Here, we examined the spatiotemporal heterogeneity in resource exploitation of a nuisance and overpopulated gull species with a great feeding plasticity, the Yellow‐legged gull Larus michahellis, by measuring the stable isotope ratios of carbon (¹³C/¹²C, δ¹³C), nitrogen (¹⁵N/¹⁴N, δ¹⁵N) and sulphur (³⁴S/³³S, δ³⁴S) in different wing feathers. Results Tracing isotopic changes among feathers in relation to moulting sequence, we showed that isotopic analyses on first primary feathers were good indicators of breeding trophic ecology, while others, such as eighth secondary feathers, reflected the feeding behaviour during the non‐breeding period. This knowledge was then used to report on estimations of seasonal feeding patterns throughout the year for seven gull populations along the Western Mediterranean coast. Main conclusions The high diversity in the exploitation patterns of the foraging habitats found in the study area both at spatial and at temporal scale indicated the opportunistic feeding behaviour of Yellow‐legged gulls. Proposed isotopic approach has revealed as a useful tool to evaluate spatiotemporal variations in the feeding ecology of populations which may become clue for dealing with the management of wild species, not only nuisance populations but also endangered species. We finally pointed out the relevance of the isotopic variability among individuals to infer diet diversity and food availability of a given population, thereby allowing demographic forecasts when trophic resources vary in abundance.