Michelle C. Mack

Consequences of changing biodiversity

Human alteration of the global environment has triggered the sixth major extinction event in the history of life and caused widespread changes in the global distribution of organisms. These changes in biodiversity alter ecosystem processes and change the resilience of ecosystems to environmental change. This has profound consequences for services that humans derive from ecosystems. The large ecological and societal consequences of changing biodiversity should be minimized to preserve options for future solutions to global environmental problems.

Ecosystem carbon storage in arctic tundra reduced by long-term nutrient fertilization

Global warming is predicted to be most pronounced at high latitudes, and observational evidence over the past 25 years suggests that this warming is already under way. One-third of the global soil carbon pool is stored in northern latitudes, so there is considerable interest in understanding how the carbon balance of northern ecosystems will respond to climate warming. Observations of controls over plant productivity in tundra and boreal ecosystems have been used to build a conceptual model of response to warming, where warmer soils and increased decomposition of plant litter increase nutrient availability, which, in turn, stimulates plant production and increases ecosystem carbon storage. Here we present the results of a long-term fertilization experiment in Alaskan tundra, in which increased nutrient availability caused a net ecosystem loss of almost 2,000 grams of carbon per square meter over 20 years. We found that annual aboveground plant production doubled during the experiment. Losses of carbon and nitrogen from deep soil layers, however, were substantial and more than offset the increased carbon and nitrogen storage in plant biomass and litter. Our study suggests that projected release of soil nutrients associated with high-latitude warming may further amplify carbon release from soils, causing a net loss of ecosystem carbon and a positive feedback to climate warming.

The Impact of Boreal Forest Fire on Climate Warming

We report measurements and analysis of a boreal forest fire, integrating the effects of greenhouse gases, aerosols, black carbon deposition on snow and sea ice, and postfire changes in surface albedo. The net effect of all agents was to increase radiative forcing during the first year (34 ± 31 Watts per square meter of burned area), but to decrease radiative forcing when averaged over an 80-year fire cycle (–2.3 ± 2.2 Watts per square meter) because multidecadal increases in surface albedo had a larger impact than fire-emitted greenhouse gases. This result implies that future increases in boreal fire may not accelerate climate warming.

Global patterns of foliar nitrogen isotopes and their relationships with climate, mycorrhizal fungi, foliar nutrient concentrations, and nitrogen availability.

Ratios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (delta(15)N), foliar N concentrations, mycorrhizal type and climate for over 11,000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar delta(15)N by 2 per thousand, 3.2 per thousand, 5.9 per thousand, respectively, relative to nonmycorrhizal plants. Foliar delta(15)N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT >or= -0.5 degrees C, but was invariant with MAT across sites with MAT < -0.5 degrees C. In independent landscape-level to regional-level studies, foliar delta(15)N increased with increasing N availability; at the global scale, foliar delta(15)N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar delta(15)N and ultimately global patterns in N cycling.

The human dimension of fire regimes on Earth

Humans and their ancestors are unique in being a fire-making species, but ‘natural’ (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human-set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from ‘natural’ background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke-related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas-induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre-human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

Carbon loss from an unprecedented Arctic tundra wildfire

Arctic tundra soils store large amounts of carbon (C) in organic soil layers hundreds to thousands of years old that insulate, and in some cases maintain, permafrost soils. Fire has been largely absent from most of this biome since the early Holocene epoch, but its frequency and extent are increasing, probably in response to climate warming. The effect of fires on the C balance of tundra landscapes, however, remains largely unknown. The Anaktuvuk River fire in 2007 burned 1,039 square kilometres of Alaska’s Arctic slope, making it the largest fire on record for the tundra biome and doubling the cumulative area burned since 1950 (ref. 5). Here we report that tundra ecosystems lost 2,016 ± 435 g C m−2 in the fire, an amount two orders of magnitude larger than annual net C exchange in undisturbed tundra. Sixty per cent of this C loss was from soil organic matter, and radiocarbon dating of residual soil layers revealed that the maximum age of soil C lost was 50 years. Scaled to the entire burned area, the fire released approximately 2.1 teragrams of C to the atmosphere, an amount similar in magnitude to the annual net C sink for the entire Arctic tundra biome averaged over the last quarter of the twentieth century. The magnitude of ecosystem C lost by fire, relative to both ecosystem and biome-scale fluxes, demonstrates that a climate-driven increase in tundra fire disturbance may represent a positive feedback, potentially offsetting Arctic greening and influencing the net C balance of the tundra biome.

The Representation of Arctic Soils in the Land Surface Model: The Importance of Mosses

Mosses dominate the surface cover in high northern latitudes and have the potential to play a key role in modifying the thermal and hydrologic regime of Arctic soils. These modifications in turn feed back to influence surface energy exchanges and hence may affect regional climate. However, mosses are poorly represented in models of the land surface. In this study the NCAR Land Surface Model (LSM) was modified in two ways. First, additional soil texture types including mosses and lichens were added to more realistically represent northern soils. Second, the LSM was also modified so that a different soil texture type could be specified for each layer. Several experiments were performed using climate data from an Arctic tundra site in 1995. The model was run for a homogeneous loam soil column and then also for columns that included moss, lichen, peat, and sand. The addition of a surface layer of moss underlain by peat and loam had a substantial impact on modeled surface processes. First, moss acted as an insulative layer producing cooler summer temperatures (6.9 8C lower at 0.5 m) and warmer winter temperatures (2.38C higher at 0.5 m) when compared with a homogenous loam soil column. Second, a soil column with a moss surface had a greater surface infiltration, leading to greater storage of soil moisture in lower layers when compared with a homogeneous loam column. Last, moss modulated the surface energy exchanges by decreasing soil heat flux (57% in July) and increasing turbulent fluxes of heat (67% in July) and moisture (15% in July). Mosses were also more effective contributors to total latent heating than was a bare loam surface. These results suggest that the addition of moss and the ability to prescribe different soil textures for different soil layers result in a more plausible distribution of heat and water within the column and that these modifications should be incorporated into regional and global climate models.

The effect of nutrient deposition on bacterial communities in Arctic tundra soil.

The microbial communities of high-latitude ecosystems are expected to experience rapid changes over the next century due to climate warming and increased deposition of reactive nitrogen, changes that will likely affect microbial community structure and function. In moist acidic tundra (MAT) soils on the North Slope of the Brooks Range, Alaska, substantial losses of C and N were previously observed after long-term nutrient additions. To analyse the role of microbial communities in these losses, we utilized 16S rRNA gene tag pyrosequencing coupled with community-level physiological profiling to describe changes in MAT bacterial communities after short- and long-term nutrient fertilization in four sets of paired control and fertilized MAT soil samples. Bacterial diversity was lower in long-term fertilized plots. The Acidobacteria were one of the most abundant phyla in all soils and distinct differences were noted in the distributions of Acidobacteria subgroups between mineral and organic soil layers that were also affected by fertilization. In addition, Alpha- and Gammaproteobacteria were more abundant in long-term fertilized samples compared with control soils. The dramatic increase in sequences within the Gammaproteobacteria identified as Dyella spp. (order Xanthomonadales) in the long-term fertilized samples was confirmed by quantitative PCR (qPCR) in several samples. Long-term fertilization was also correlated with shifts in the utilization of specific substrates by microbes present in the soils. The combined data indicate that long-term fertilization resulted in a significant change in microbial community structure and function linked to changes in carbon and nitrogen availability and shifts in above-ground plant communities.

Plant Species Composition and Productivity following Permafrost Thaw and Thermokarst in Alaskan Tundra

Climate warming is expected to have a large impact on plant species composition and productivity in northern latitude ecosystems. Warming can affect vegetation communities directly through temperature effects on plant growth and indirectly through alteration of soil nutrient availability. In addition, warming can cause permafrost to thaw and thermokarst (ground subsidence) to develop, which can alter the structure of the ecosystem by altering hydrological patterns within a site. These multiple direct and indirect effects of permafrost thawing are difficult to simulate in experimental approaches that often manipulate only one or two factors. Here, we used a natural gradient approach with three sites to represent stages in the process of permafrost thawing and thermokarst. We found that vascular plant biomass shifted from graminoid-dominated tundra in the least disturbed site to shrub-dominated tundra at the oldest, most subsided site, whereas the intermediate site was co-dominated by both plant functional groups. Vascular plant productivity patterns followed the changes in biomass, whereas nonvascular moss productivity was especially important in the oldest, most subsided site. The coefficient of variation for soil moisture was higher in the oldest, most subsided site suggesting that in addition to more wet microsites, there were other microsites that were drier. Across all sites, graminoids preferred the cold, dry microsites whereas the moss and shrubs were associated with the warm, moist microsites. Total nitrogen contained in green plant biomass differed across sites, suggesting that there were increases in soil nitrogen availability where permafrost had thawed.

Postfire Soil N Cycling in Northern Conifer Forests Affected by Severe, Stand-Replacing Wildfires

Severe, stand-replacing fires affect large areas of northern temperate and boreal forests, potentially modifying ecosystem function for decades after their occurrence. Because these fires occur over large extents, and in areas where plant production is limited by nitrogen (N) availability, the effect of fire on N cycling may be important for long-term ecosystem productivity. In this article, we review what is known about postfire N cycling in northern temperate and boreal forests experiencing stand-replacing fires. We then build upon existing literature to identify the most important mechanisms that control postfire N availability in systems experiencing severe, stand-replacing fires compared with fires of lower severity. These mechanisms include changes in abiotic conditions caused by the opening of the canopy (for example, decreased LAI, increased solar radiation), changes in ground layer quantity and quality (for example, nutrient release, permafrost levels), and postfire plant and microbial adaptations affecting N fixation and N uptake (for example, serotiny, germination cues). Based on the available literature, these mechanisms appear to affect N inputs, internal N cycling, and N outputs in various ways, indicating that severe fire systems are variable across time and space as a result of complex interactions between postfire abiotic and biotic factors. Future experimental work should be focused on understanding these mechanisms and their variability across the landscape.