Convergent ecosystem responses to 23‐year ambient and manipulated warming link advancing snowmelt and shrub encroachment to transient and long‐term climate–soil carbon feedback

Ecosystem responses to climate change can exert positive or negative feedbacks on climate, mediated in part by slow‐moving factors such as shifts in vegetation community composition. Long‐term experimental manipulations can be used to examine such ecosystem responses, but they also present another opportunity: inferring the extent to which contemporary climate change is responsible for slow changes in ecosystems under ambient conditions. Here, using 23 years of data, we document a shift from nonwoody to woody vegetation and a loss of soil carbon in ambient plots and show that these changes track previously shown similar but faster changes under experimental warming. This allows us to infer that climate change is the cause of the observed shifts in ambient vegetation and soil carbon and that the vegetation responses mediate the observed changes in soil carbon. Our findings demonstrate the realism of an experimental manipulation, allow attribution of a climate cause to observed ambient ecosystem changes, and demonstrate how a combination of long‐term study of ambient and experimental responses to warming can identify mechanistic drivers needed for realistic predictions of the conditions under which ecosystems are likely to become carbon sources or sinks over varying timescales.     

Long‐term carbon storage through retention of dissolved aromatic acids by reactive particles in soil

Soils retain large quantities of carbon, thereby slowing its return to the atmosphere. The mechanisms governing organic carbon sequestration in soil remain poorly understood, yet are integral to understanding soil‐climate feedbacks. We evaluated the biochemistry of dissolved and solid organic carbon in potential source and sink horizons across a chronosequence of volcanic soils in Hawai'i. The soils are derived from similar basaltic parent material on gently sloping volcanic shield surfaces, support the same vegetation assemblage, and yet exhibit strong shifts in soil mineralogy and soil carbon content as a function of volcanic substrate age. Solid‐state¹³carbon nuclear magnetic resonance spectra indicate that the most persistent mineral‐bound carbon is comprised of partially oxidized aromatic compounds with strong chemical resemblance to dissolved organic matter derived from plant litter. A molecular mixing model indicates that protein, lipid, carbohydrate, and char content decreased whereas oxidized lignin and carboxyl/carbonyl content increased with increasing short‐range order mineral content. When solutions rich in dissolved organic matter were passed through Bw‐horizon mineral cores, aromatic compounds were preferentially sorbed with the greatest retention occurring in horizons containing the greatest amount of short‐range ordered minerals. These minerals are reactive metastable nanocrystals that are most common in volcanic soils, but exist in smaller amounts in nearly all major soil classes. Our results indicate that long‐term carbon storage in short‐range ordered minerals occurs via chemical retention with dissolved aromatic acids derived from plant litter and carried along preferential flow‐paths to deeper B horizons.     

Long‐term antagonistic effect of increased precipitation and nitrogen addition on soil respiration in a semiarid steppe

Changes in water and nitrogen (N) availability due to climate change and atmospheric N deposition could have significant effects on soil respiration, a major pathway of carbon (C) loss from terrestrial ecosystems. A manipulative experiment simulating increased precipitation and atmospheric N deposition has been conducted for 9 years (2005–2013) in a semiarid grassland in Mongolian Plateau, China. Increased precipitation and N addition interactively affect soil respiration through the 9 years. The interactions demonstrated that N addition weakened the precipitation‐induced stimulation of soil respiration, whereas increased precipitation exacerbated the negative impacts of N addition. The main effects of increased precipitation and N addition treatment on soil respiration were 15.8% stimulated and 14.2% suppressed, respectively. Moreover, a declining pattern and 2‐year oscillation were observed for soil respiration response to N addition under increased precipitation. The dependence of soil respiration upon gross primary productivity and soil moisture, but not soil temperature, suggests that resources C substrate supply and water availability are more important than temperature in regulating interannual variations of soil C release in semiarid grassland ecosystems. The findings indicate that atmospheric N deposition may have the potential to mitigate soil C loss induced by increased precipitation, and highlight that long‐term and multi‐factor global change studies are critical for predicting the general patterns of terrestrial C cycling in response to global change in the future.     

Short‐term carbon cycling responses of a mature eucalypt woodland to gradual stepwise enrichment of atmospheric CO2 concentration

Projections of future climate are highly sensitive to uncertainties regarding carbon (C) uptake and storage by terrestrial ecosystems. The Eucalyptus Free‐Air CO₂ Enrichment (EucFACE) experiment was established to study the effects of elevated atmospheric CO₂ concentrations (eCO₂) on a native mature eucalypt woodland with low fertility soils in southeast Australia. In contrast to other FACE experiments, the concentration of CO₂ at EucFACE was increased gradually in steps above ambient (+0, 30, 60, 90, 120, and 150 ppm CO₂ above ambient of ~400 ppm), with each step lasting approximately 5 weeks. This provided a unique opportunity to study the short‐term (weeks to months) response of C cycle flux components to eCO₂ across a range of CO₂ concentrations in an intact ecosystem. Soil CO₂ efflux (i.e., soil respiration or R_soil) increased in response to initial enrichment (e.g., +30 and +60 ppm CO₂) but did not continue to increase as the CO₂ enrichment was stepped up to higher concentrations. Light‐saturated photosynthesis of canopy leaves (A_sat) also showed similar stimulation by elevated CO₂ at +60 ppm as at +150 ppm CO₂. The lack of significant effects of eCO₂ on soil moisture, microbial biomass, or activity suggests that the increase in R_soil likely reflected increased root and rhizosphere respiration rather than increased microbial decomposition of soil organic matter. This rapid increase in R_soil suggests that under eCO_2, additional photosynthate was produced, transported belowground, and respired. The consequences of this increased belowground activity and whether it is sustained through time in mature ecosystems under eCO₂ are a priority for future research.     

Impacts of multiple extreme winter warming events on sub‐Arctic heathland: phenology,reproduction, growth,and CO2 flux responses

Extreme weather events can have strong negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme, short‐lived, winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (for instance, 2–10 °C for 2–14 days) but upon return to normal winter climate exposes the ecosystem to much colder temperatures due to the loss of insulating snow. Single events have been shown to reduce plant reproduction and increase shoot mortality, but impacts of multiple events are little understood as are the broader impacts on community structure, growth, carbon balance, and nutrient cycling. To address these issues, we simulated week‐long extreme winter warming events – using infrared heating lamps and soil warming cables – for 3 consecutive years in a sub‐Arctic heathland dominated by the dwarf shrubs Empetrum hermaphroditum, Vaccinium vitis‐idaea (both evergreen) and Vaccinium myrtillus (deciduous). During the growing seasons after the second and third winter event, spring bud burst was delayed by up to a week for E. hermaphroditum and V. myrtillus, and berry production reduced by 11–75% and 52–95% for E. hermaphroditum and V. myrtillus, respectively. Greater shoot mortality occurred in E. hermaphroditum (up to 52%), V. vitis‐idaea (51%), and V. myrtillus (80%). Root growth was reduced by more than 25% but soil nutrient availability remained unaffected. Gross primary productivity was reduced by more than 50% in the summer following the third simulation. Overall, the extent of damage was considerable, and critically plant responses were opposite in direction to the increased growth seen in long‐term summer warming simulations and the ‘greening’ seen for some arctic regions. Given the Arctic is warming more in winter than summer, and extreme events are predicted to become more frequent, this generates large uncertainty in our current understanding of arctic ecosystem responses to climate change.     

Long‐term enhanced winter soil frost alters growing season CO2 fluxes through its impact on vegetation development in a boreal peatland

At high latitudes, winter climate change alters snow cover and, consequently, may cause a sustained change in soil frost dynamics. Altered winter soil conditions could influence the ecosystem exchange of carbon dioxide (CO₂) and, in turn, provide feedbacks to ongoing climate change. To investigate the mechanisms that modify the peatland CO₂ exchange in response to altered winter soil frost, we conducted a snow exclusion experiment to enhance winter soil frost and to evaluate its short‐term (1–3 years) and long‐term (11 years) effects on CO₂ fluxes during subsequent growing seasons in a boreal peatland. In the first 3 years after initiating the treatment, no significant effects were observed on either gross primary production (GPP) or ecosystem respiration (ER). However, after 11 years, the temperature sensitivity of ER was reduced in the treatment plots relative to the control, resulting in an overall lower ER in the former. Furthermore, early growing season GPP was also lower in the treatment plots than in the controls during periods with photosynthetic photon flux density (PPFD) ≥800 μmol m^?2 s^?1, corresponding to lower sedge leaf biomass in the treatment plots during the same period. During the peak growing season, a higher GPP was observed in the treatment plots under the low light condition (i.e. PPFD 400 μmol m^?2 s^?1) compared to the control. As Sphagnum moss maximizes photosynthesis at low light levels, this GPP difference between the plots may have been due to greater moss photosynthesis, as indicated by greater moss biomass production, in the treatment plots relative to the controls. Our study highlights the different responses to enhanced winter soil frost among plant functional types which regulate CO₂ fluxes, suggesting that winter climate change could considerably alter the growing season CO₂ exchange in boreal peatlands through its effect on vegetation development.     

Does deep soil N availability sustain long‐term ecosystem responses to elevated CO2?

A scrub‐oak woodland has maintained higher aboveground biomass accumulation after 11 years of atmospheric CO₂ enrichment (ambient +350 μmol CO₂ mol^?1), despite the expectation of strong nitrogen (N) limitation at the site. We hypothesized that changes in plant available N and exploitation of deep sources of inorganic N in soils have sustained greater growth at elevated CO₂. We employed a suite of assays performed in the sixth and 11th year of a CO₂ enrichment experiment designed to assess soil N dynamics and N availability in the entire soil profile. In the 11th year, we found no differences in gross N flux, but significantly greater microbial respiration (P≤0.01) at elevated CO₂. Elevated CO₂ lowered extractable inorganic N concentrations (P=0.096) considering the whole soil profile (0–190 cm). Conversely, potential net N mineralization, although not significant in considering the entire profile (P=0.460), tended to be greater at elevated CO₂. Ion‐exchange resins placed in the soil profile for approximately 1 year revealed that potential N availability at the water table was almost 3 × greater than found elsewhere in the profile, and we found direct evidence using a ¹⁵N tracer study that plants took up N from the water table. Increased microbial respiration and shorter mean residence times of inorganic N at shallower depths suggests that enhanced SOM decomposition may promote a sustained supply of inorganic N at elevated CO₂. Deep soil N availability at the water table is considerable, and provides a readily available source of N for plant uptake. Increased plant growth at elevated CO₂ in this ecosystem may be sustained through greater inorganic N supply from shallow soils and N uptake from deep soil.     

Long‐term evolutionary and ecological responses of calcifying phytoplankton to changes in atmospheric CO2

Calcifying phytoplankton play an important role in marine ecosystems and global biogeochemical cycles, affecting the transfer of both organic and inorganic carbon from the surface to the deep ocean. Coccolithophores are the most prominent members of this group, being well adapted to low‐nutrients environments (e.g., subtropical gyres). Despite urgent concerns, their response to rising atmospheric carbon dioxide levels (pCO₂) and ocean acidification is still poorly understood, and short‐term experiments may not extrapolate into longer‐term climatic adaptation. Current atmospheric pCO₂ (~390 ppmv) is unprecedented since at least 3 million years ago (Ma), and levels projected for the next century were last seen more than 34 Ma. Hence, a deep‐time perspective is needed to understand the long‐term effects of high pCO₂ on the biosphere. Here we combine a comprehensive fossil data set on coccolithophore cell size with a novel measure of ecological prominence: Summed Common Species Occurrence Rate (SCOR). The SCOR is decoupled from species richness, and captures changes in the extent to which coccolithophores were common and widespread, based on global occurrences in deep‐sea sediments. The size and SCOR records are compared to state‐of‐the‐art data on climatic and environmental changes from 50 to 5 Ma. We advance beyond simple correlations and trends to quantify the relative strength and directionality of information transfer among these records. Coccolithophores were globally more common and widespread, larger, and more heavily calcified in the pre‐34 Ma greenhouse world, and declined along with pCO₂ during the Oligocene (34–23 Ma). Our results suggest that atmospheric pCO₂ has exerted an important long‐term control on coccolithophores, directly through its availability for photosynthesis or indirectly via weathering supply of resources for growth and calcification.     

Long‐term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil

Nitrogen (N) deposition is a component of global change that has considerable impact on belowground carbon (C) dynamics. Plant growth stimulation and alterations of fungal community composition and functions are the main mechanisms driving soil C gains following N deposition in N‐limited temperate forests. In N‐rich tropical forests, however, N deposition generally has minor effects on plant growth; consequently, C storage in soil may strongly depend on the microbial processes that drive litter and soil organic matter decomposition. Here, we investigated how microbial functions in old‐growth tropical forest soil responded to 13 years of N addition at four rates: 0 (Control), 50 (Low‐N), 100 (Medium‐N), and 150 (High‐N) kg N ha^?1 year^?1. Soil organic carbon (SOC) content increased under High‐N, corresponding to a 33% decrease in CO₂ efflux, and reductions in relative abundances of bacteria as well as genes responsible for cellulose and chitin degradation. A 113% increase in N₂O emission was positively correlated with soil acidification and an increase in the relative abundances of denitrification genes (narG and norB). Soil acidification induced by N addition decreased available P concentrations, and was associated with reductions in the relative abundance of phytase. The decreased relative abundance of bacteria and key functional gene groups for C degradation were related to slower SOC decomposition, indicating the key mechanisms driving SOC accumulation in the tropical forest soil subjected to High‐N addition. However, changes in microbial functional groups associated with N and P cycling led to coincidentally large increases in N₂O emissions, and exacerbated soil P deficiency. These two factors partially offset the perceived beneficial effects of N addition on SOC storage in tropical forest soils. These findings suggest a potential to incorporate microbial community and functions into Earth system models considering their effects on greenhouse gas emission, biogeochemical processes, and biodiversity of tropical ecosystems.     

Long‐term terrestrial carbon dynamics in the Midwestern United States during 1850–2015: Roles of land use and cover change and agricultural management

To meet the increasing food and biofuel demand, the Midwestern United States has become one of the most intensively human‐disturbed hotspots, characterized by widespread cropland expansion and various management practices. However, the role of human activities in the carbon (C) cycling across managed landscape remains far from certain. In this study, based on state‐ and national census, field experiments, and model simulation, we comprehensively examined long‐term carbon storage change in response to land use and cover change (LUCC) and agricultural management in the Midwest from 1850 to 2015. We also quantified estimation uncertainties related to key parameter values. Model estimation showed LUCC led to a reduction of 1.35 Pg (with a range of 1.3–1.4 Pg) in vegetation C pool of the Midwest, yet agricultural management barely affected vegetation C change. In comparison, LUCC reduced SOC by 4.5 Pg (3.1 to 6.2 Pg), while agricultural management practices increased SOC stock by 0.9 Pg. Moreover, we found 45% of the study area was characterized by continuously decreasing SOC caused by LUCC, and SOC in 13% and 31% of the area was fully and partially recovered, respectively, since 1850. Agricultural management was estimated to increase the area of full recovery and partial recovery by 8.5% and 1.1%. Our results imply that LUCC plays an essential role in regional C balance, and more importantly, sustainable land management can be beneficial for strengthening C sequestration of the agroecosystems in the Midwestern US, which may serve as an important contributor to C sinks in the US.     

Simulating effects of changing climate and CO2 emissions on soil carbon pools at the Hubbard Brook experimental forest

Carbon (C) sequestration in forest biomass and soils may help decrease regional C footprints and mitigate future climate change. The efficacy of these practices must be verified by monitoring and by approved calculation methods (i.e., models) to be credible in C markets. Two widely used soil organic matter models – CENTURY and RothC – were used to project changes in SOC pools after clear‐cutting disturbance, as well as under a range of future climate and atmospheric carbon dioxide (CO₂) scenarios. Data from the temperate, predominantly deciduous Hubbard Brook Experimental Forest (HBEF) in New Hampshire, USA, were used to parameterize and validate the models. Clear‐cutting simulations demonstrated that both models can effectively simulate soil C dynamics in the northern hardwood forest when adequately parameterized. The minimum postharvest SOC predicted by RothC occurred in postharvest year 14 and was within 1.5% of the observed minimum, which occurred in year 8. CENTURY predicted the postharvest minimum SOC to occur in year 45, at a value 6.9% greater than the observed minimum; the slow response of both models to disturbance suggests that they may overestimate the time required to reach new steady‐state conditions. Four climate change scenarios were used to simulate future changes in SOC pools. Climate‐change simulations predicted increases in SOC by as much as 7% at the end of this century, partially offsetting future CO₂ emissions. This sequestration was the product of enhanced forest productivity, and associated litter input to the soil, due to increased temperature, precipitation and CO₂. The simulations also suggested that considerable losses of SOC (8–30%) could occur if forest vegetation at HBEF does not respond to changes in climate and CO₂ levels. Therefore, the source/sink behavior of temperate forest soils likely depends on the degree to which forest growth is stimulated by new climate and CO₂ conditions.     

The effect of fire and permafrost interactions on soil carbon accumulation in an upland black spruce ecosystem of interior Alaska: implications for post‐thaw carbon loss

High‐latitude regions store large amounts of organic carbon (OC) in active‐layer soils and permafrost, accounting for nearly half of the global belowground OC pool. In the boreal region, recent warming has promoted changes in the fire regime, which may exacerbate rates of permafrost thaw and alter soil OC dynamics in both organic and mineral soil. We examined how interactions between fire and permafrost govern rates of soil OC accumulation in organic horizons, mineral soil of the active layer, and near‐surface permafrost in a black spruce ecosystem of interior Alaska. To estimate OC accumulation rates, we used chronosequence, radiocarbon, and modeling approaches. We also developed a simple model to track long‐term changes in soil OC stocks over past fire cycles and to evaluate the response of OC stocks to future changes in the fire regime. Our chronosequence and radiocarbon data indicate that OC turnover varies with soil depth, with fastest turnover occurring in shallow organic horizons (～60 years) and slowest turnover in near‐surface permafrost (>3000 years). Modeling analysis indicates that OC accumulation in organic horizons was strongly governed by carbon losses via combustion and burial of charred remains in deep organic horizons. OC accumulation in mineral soil was influenced by active layer depth, which determined the proportion of mineral OC in a thawed or frozen state and thus, determined loss rates via decomposition. Our model results suggest that future changes in fire regime will result in substantial reductions in OC stocks, largely from the deep organic horizon. Additional OC losses will result from fire‐induced thawing of near‐surface permafrost. From these findings, we conclude that the vulnerability of deep OC stocks to future warming is closely linked to the sensitivity of permafrost to wildfire disturbance.     

Long‐term increases in tropical flowering activity across growth forms in response to rising CO2 and climate change

Mounting evidence suggests that anthropogenic global change is altering plant species composition in tropical forests. Fewer studies, however, have focused on long‐term trends in reproductive activity, in part because of the lack of data from tropical sites. Here, we analyze a 28‐year record of tropical flower phenology in response to anthropogenic climate and atmospheric change. We show that a multidecadal increase in flower activity is most strongly associated with rising atmospheric CO₂ concentrations using yearly aggregated data. Compared to significant climatic factors, CO₂ had on average an approximately three‐, four‐, or fivefold stronger effect than rainfall, solar radiation, and the Multivariate ENSO Index, respectively. Peaks in flower activity were associated with greater solar radiation and lower rainfall during El Niño years. The effect of atmospheric CO₂ on flowering has diminished over the most recent decade for lianas and canopy trees, whereas flowering of midstory trees and shrub species continued to increase with rising CO₂. Increases in flowering were accompanied by a lengthening of flowering duration for canopy and midstory trees. Understory treelets did not show increases in flowering but did show increases in duration. Given that atmospheric CO₂ will likely continue to climb over the next century, a long‐term increase in flowering activity may persist in some growth forms until checked by nutrient limitation or by climate change through rising temperatures, increasing drought frequency and/or increasing cloudiness and reduced insolation.