Permafrost thaw and the subsequent mobilization of carbon (C) stored in previously frozen soil organic matter (SOM) have the potential to be a strong positive feedback to climate. As the northern ...permafrost region experiences as much as a doubling of the rate of warming as the rest of the Earth, the vast amount of C in permafrost soils is vulnerable to thaw, decomposition and release as atmospheric greenhouse gases. Diagnostic and predictive estimates of high-latitude terrestrial C fluxes vary widely among different models depending on how dynamics in permafrost, and the seasonally thawed 'active layer' above it, are represented. Here, we employ a process-based model simulation experiment to assess the net effect of active layer dynamics on this 'permafrost carbon feedback' in recent decades, from 1970 to 2006, over the circumpolar domain of continuous and discontinuous permafrost. Over this time period, the model estimates a mean increase of 6.8 cm in active layer thickness across the domain, which exposes a total of 11.6 Pg C of thawed SOM to decomposition. According to our simulation experiment, mobilization of this previously frozen C results in an estimated cumulative net source of 3.7 Pg C to the atmosphere since 1970 directly tied to active layer dynamics. Enhanced decomposition from the newly exposed SOM accounts for the release of both CO2 (4.0 Pg C) and CH4 (0.03 Pg C), but is partially compensated by CO2 uptake (0.3 Pg C) associated with enhanced net primary production of vegetation. This estimated net C transfer to the atmosphere from permafrost thaw represents a significant factor in the overall ecosystem carbon budget of the Pan-Arctic, and a non-trivial additional contribution on top of the combined fossil fuel emissions from the eight Arctic nations over this time period.
Studies indicate that, historically, terrestrial ecosystems of the northern high-latitude region may have been responsible for up to 60% of the global net land-based sink for atmospheric CO sub(2). ...However, these regions have recently experienced remarkable modification of the major driving forces of the carbon cycle, including surface air temperature warming that is significantly greater than the global average and associated increases in the frequency and severity of disturbances. Whether Arctic tundra and boreal forest ecosystems will continue to sequester atmospheric CO sub(2) in the face of these dramatic changes is unknown. Here we show the results of model simulations that estimate a 41 Tg C yr super(-1) sink in the boreal land regions from 1997 to 2006, which represents a 73% reduction in the strength of the sink estimated for previous decades in the late 20th century. Our results suggest that CO sub(2) uptake by the region in previous decades may not be as strong as previously estimated. The recent decline in sink strength is the combined result of (1) weakening sinks due to warming-induced increases in soil organic matter decomposition and (2) strengthening sources from pyrogenic CO sub(2) emissions as a result of the substantial area of boreal forest burned in wildfires across the region in recent years. Such changes create positive feedbacks to the climate system that accelerate global warming, putting further pressure on emission reductions to achieve atmospheric stabilization targets. Key Points * Reduction in strength of the boreal land CO2 sink * Sink strength not as strong as previously estimated * Weakening sink due to climate change and disturbance
We use the Terrestrial Ecosystem Model (TEM, Version 4.1) and the land cover data set of the international geosphere'biosphere program to investigate how increasing atmospheric CO
2
concentration and ...climate variability during 1900-1994 affect the carbon storage of terrestrial ecosystems in the conterminous USA, and how carbon storage has been affected by land-use change. The estimates of TEM indicate that over the past 95 years a combination of increasing atmospheric CO
2
with historical temperature and precipitation variability causes a 4.2% (4.3 Pg C) decrease in total carbon storage of potential vegetation in the conterminous US, with vegetation carbon decreasing by 7.2% (3.2 Pg C) and soil organic carbon decreasing by 1.9% (1.1 Pg C). Several dry periods including the 1930s and 1950s are responsible for the loss of carbon storage. Our factorial experiments indicate that precipitation variability alone decreases total carbon storage by 9.5%. Temperature variability alone does not significantly affect carbon storage. The effect of CO
2
fertilization alone increases total carbon storage by 4.4%. The effects of increasing atmospheric CO
2
and climate variability are not additive. Interactions among CO
2
, temperature and precipitation increase total carbon storage by 1.1%. Our study also shows substantial year-to-year variations in net carbon exchange between the atmosphere and terrestrial ecosystems due to climate variability. Since the 1960s, we estimate these terrestrial ecosystems have acted primarily as a sink of atmospheric CO
2
as a result of wetter weather and higher atmospheric CO
2
concentrations. For the 1980s, we estimate the natural terrestrial ecosystems, excluding cropland and urban areas, of the conterminous US have accumulated 78.2 Tg C yr
-1
because of the combined effect of increasing atmospheric CO
2
and climate variability. For the conterminous US, we estimate that the conversion of natural ecosystems to cropland and urban areas has caused a 18.2% (17.7 Pg C) reduction in total carbon storage from that estimated for potential vegetation. The carbon sink capacity of natural terrestrial ecosystems in the conterminous US is about 69% of that estimated for potential vegetation.
Because model predictions at continental and global scales are necessarily based on broad characterizations of vegetation, soils, and climate, estimates of carbon stocks and fluxes made by global ...terrestrial biosphere models may not be accurate for every region. At the regional scale, we suggest that attention can be focused more clearly on understanding the relative strengths of predicted net primary productivity (NPP) limitation by energy, water, and nutrients. We evaluate the sources of variability among model predictions of NPP with a regional-scale comparison between estimates made by PnET-II (a forest ecosystem process model previously applied to the northeastern region) and TEM 4.0 (a terrestrial biosphere model typically applied to the globe) for the northeastern US. When the same climate, vegetation, and soil data sets were used to drive both models, regional average NPP predictions made by PnET-II and TEM were remarkably similar, and at the biome level, model predictions agreed fairly well with NPP estimates developed from field measurements. However, TEM 4.0 predictions were more sensitive to regional variations in temperature as a result of feedbacks between temperature and belowground N availability. In PnET-II, the direct link between transpiration and photosynthesis caused substantial water stress in hardwood and pine forest types with increases in solar radiation; predicted water stress was relieved substantially when soil water holding capacity (WHC) was increased. Increasing soil WHC had little effect on TEM 4.0 predictions because soil water storage was already sufficient to meet plant demand with baseline WHC values, and because predicted N availability under baseline conditions in this region was not limited by water. Because NPP predictions were closely keyed to forest cover type, the relative coverage of low- versus high-productivity forests at both fine and coarse resolutions was an important determinant of regional NPP predictions. Therefore, changes in grid cell size and differences in the methods used to aggregate from fine to coarse resolution were important to NPP predictions insofar as they changed the relative proportions of forest cover. We suggest that because the small patches of high-elevation spruce-fir forest in this region are substantially less productive than forests in the remainder of the region, more accurate NPP predictions will result if models applied to this region use land cover input data sets that retain as much fine-resolution forest type variability as possible. The differences among model responses to variations in climate and soil WHC data sets suggest that the models will respond quite differently to scenarios of future climate. A better understanding of the dynamic interactions between water stress, N availability, and forest productivity in this region will enable models to make more accurate predictions of future carbon stocks and fluxes.
We use the Multiple Element Limitation (MEL) model to examine responses of 12 ecosystems to elevated carbon dioxide (CO2), warming, and 20% decreases or increases in precipitation. Ecosystems respond ...synergistically to elevated CO2, warming, and decreased precipitation combined because higher water‐use efficiency with elevated CO2 and higher fertility with warming compensate for responses to drought. Response to elevated CO2, warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO2 and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C‐nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO2 and climate change.
ABSTRACT
We used the terrestrial ecosystem model (TEM, version 4.0) to estimate global responses of annual net primary production (NPP) and total carbon storage to changes in climate and atmospheric ...CO2, driven by the climate outputs from the 2‐dimensional MIT L‐O climate model and the 3‐dimensional GISS and GFDL‐q atmospheric general circulation models (GCMs). For contemporary climate with 315 ppmv CO2, TEM estimates that global NPP is 47.9 PgC/ yr and global total carbon storage is 1658 PgC: 908 PgC of vegetation carbon and 750 PgC of reactive soil organic carbon. For climate change associated with a doubling of radiative forcing and an atmospheric level of 522 ppmv CO2, the responses of global NPP are + 17.8% for the MIT L‐O climate, + 18.5% for the GFDL‐q climate and + 20.6% for the GISS climate. The responses of global total carbon storage are + 6.9% for the MIT L‐O climate, + 8.3% for GFDL‐q climate and + 8.7% for the GISS climate. Among the three climate change predictions, the changes in latitudinal distributions of cumulative NPP and total carbon storage along 0.5° latitudinal bands vary slightly, except in high latitudes. There are generally minor differences in cumulative NPP and total carbon storage for most of the 18 biomes, except for the responses of total carbon storage in boreal biomes for the 2‐D MIT L‐O climate change. The results demonstrate that the linkage between the TEM and the 2‐D climate model is useful for impact assessment and uncertainty analysis within an integrated assessment framework at the scales of the globe, economic regions and biomes, given the compromise between computational efficiency in the 2‐D climate model and more detailed spatial representation of climate fields in 3‐D GCMs.
Molecules of keystone significance are relatively rare, yet mediate a variety of interactions between organisms. They influence the distribution and abundance of species, the transfer of energy ...across multiple trophic levels, and thus they play significant roles in structuring ecosystems. Despite their potential importance in facilitating our understanding of ecological systems, only three molecules thus far have been proposed as molecules of keystone significance: saxitoxin and dimethyl sulfide in marine communities and tetrodotoxin in riparian communities. In the course of studying the neuroecology of chemical defenses, we identified three mycosporine-like amino acids (MAAs)--N-ethanol palythine (= asterina-330), N-isopropanol palythine (= aplysiapalythine A), and N-ethyl palythine (= aplysiapalythine B)--as intraspecific alarm cues for sea hares (Aplysia californica). These alarm cues are released in the ink secretion of sea hares and cause avoidance behaviors in neighboring conspecifics. Further, we show that these three bioactive MAAs, two aplysiapalythine A (APA) and -B (APB) being previously unknown molecules, are present in the algal diet of sea hares and are concentrated in their defensive secretion as well as in their skin. MAAs are known to be produced by algae, fungi, and cyanobacteria and are acquired by many aquatic animals through trophic interactions. MAAs are widely used as sunscreens, among other uses, but sea hares modify their function to serve a previously undocumented role, as intraspecific chemical cues. Our findings highlight the multifunctionality of MAAs and their role in ecological connectivity, suggesting that they may function as molecules of keystone significance in marine ecosystems.
There is substantial evidence that soil thermal dynamics are changing in terrestrial ecosystems of the Northern Hemisphere and that these dynamics have implications for the exchange of carbon between ...terrestrial ecosystems and the atmosphere. To date, large-scale biogeochemical models have been slow to incorporate the effects of soil thermal dynamics on processes that affect carbon exchange with the atmosphere. In this study we incorporated a soil thermal module (STM), appropriate to both permafrost and non-permafrost soils, into a large-scale ecosystem model, version 5.0 of the Terrestrial Ecosystem Model (TEM). We then compared observed regional and seasonal patterns of atmospheric CO
2
to simulations of carbon dynamics for terrestrial ecosystems north of 30°N between TEM 5.0 and an earlier version of TEM (version 4.2) that lacked a STM. The timing of the draw-down of atmospheric CO
2
at the start of the growing season and the degree of draw-down during the growing season were substantially improved by the consideration of soil thermal dynamics. Both versions of TEM indicate that climate variability and change promoted the loss of carbon from temperate ecosystems during the first half of the 20th century, and promoted carbon storage during the second half of the century. The results of the simulations by TEM suggest that land-use change in temperate latitudes (30-60°N) plays a stronger role than climate change in driving trends for increased uptake of carbon in extratropical terrestrial ecosystems (30-90°N) during recent decades. In the 1980s the TEM 5.0 simulation estimated that extratropical terrestrial ecosystems stored 0.55 Pg C yr
−1
, with 0.24 Pg C yr
−1
in North America and 0.31 Pg C yr
−1
in northern Eurasia. From 1990 through 1995 the model simulated that these ecosystems stored 0.90 Pg C yr
−1
, with 0.27 Pg C yr
−1
stored in North America and 0.63 Pg C yr
−1
stored in northern Eurasia. Thus, in comparison to the 1980s, simulated net carbon storage in the 1990s was enhanced by an additional 0.35 Pg C yr
−1
in extratropical terrestrial ecosystems, with most of the additional storage in northern Eurasia. The carbon storage simulated by TEM 5.0 in the 1980s and 1990s was lower than estimates based on other methodologies, including estimates by atmospheric inversion models and remote sensing and inventory analyses. This suggests that other issues besides the role of soil thermal dynamics may be responsible, in part, for the temporal and spatial dynamics of carbon storage of extratropical terrestrial ecosystems. In conclusion, the consideration of soil thermal dynamics and terrestrial cryospheric processes in modeling the global carbon cycle has helped to reduce biases in the simulation of the seasonality of carbon dynamics of extratropical terrestrial ecosystems. This progress should lead to an enhanced ability to clarify the role of other issues that influence carbon dynamics in terrestrial regions that experience seasonal freezing and thawing of soil.