Sensitivity of primary production to precipitation across the United States.

Primary production, a key regulator of the global carbon cycle, is highly responsive to variations in climate. Yet, a detailed, continental-scale risk assessment of climate-related impacts on primary production is lacking. We combined 16 years of MODIS NDVI data, a remotely sensed proxy for primary production, with observations from 1218 climate stations to derive values of ecosystem sensitivity to precipitation and aridity. For the first time, we produced an empirically-derived map of ecosystem sensitivity to climate across the conterminous United States. Over this 16-year period, annual primary production values were most sensitive to precipitation and aridity in dryland and grassland ecosystems. Century-long trends measured at the climate stations showed intensifying aridity and climatic variability in many of these sensitive regions. Dryland ecosystems in the western US may be particularly vulnerable to reductions in primary production and consequent degradation of ecosystem services as climate change and variability increase in the future.

Climate sensitivity functions and net primary production: A framework for incorporating climate mean and variability.

Understanding controls on net primary production (NPP) has been a long-standing goal in ecology. Climate is a well-known control on NPP, although the temporal differences among years within a site are often weaker than the spatial pattern of differences across sites. Climate sensitivity functions describe the relationship between an ecological response (e.g., NPP) and both the mean and variance of its climate driver (e.g., aridity index), providing a novel framework for understanding how climate trends in both mean and variance vary with NPP over time. Nonlinearities in these functions predict whether an increase in climate variance will have a positive effect (convex nonlinearity) or negative effect (concave nonlinearity) on NPP. The influence of climate variance may be particularly intense at ecosystem transition zones, if species reach physiological thresholds that create nonlinearities at these ecotones. Long-term data collected at the confluence of three dryland ecosystems in central New Mexico revealed that each ecosystem exhibited a unique climate sensitivity function that was consistent with long-term vegetation change occurring at their ecotones. Our analysis suggests that rising temperatures in drylands could alter the nonlinearities that determine the relative costs and benefits of variance in precipitation for primary production.

CO2 exchange and evapotranspiration across dryland ecosystems of southwestern North America.

Global-scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO2 sink. However, such analyses are poorly constrained by measured CO2 exchange in drylands. Here we address this observation gap with eddy covariance data from 25 sites in the water-limited Southwest region of North America with observed ranges in annual precipitation of 100-1000 mm, annual temperatures of 2-25°C, and records of 3-10 years (150 site-years in total). Annual fluxes were integrated using site-specific ecohydrologic years to group precipitation with resulting ecosystem exchanges. We found a wide range of carbon sink/source function, with mean annual net ecosystem production (NEP) varying from -350 to +330 gCm-2 across sites with diverse vegetation types, contrasting with the more constant sink typically measured in mesic ecosystems. In this region, only forest-dominated sites were consistent carbon sinks. Interannual variability of NEP, gross ecosystem production (GEP), and ecosystem respiration (Reco ) was larger than for mesic regions, and half the sites switched between functioning as C sinks/C sources in wet/dry years. The sites demonstrated coherent responses of GEP and NEP to anomalies in annual evapotranspiration (ET), used here as a proxy for annually available water after hydrologic losses. Notably, GEP and Reco were negatively related to temperature, both interannually within site and spatially across sites, in contrast to positive temperature effects commonly reported for mesic ecosystems. Models based on MODIS satellite observations matched the cross-site spatial pattern in mean annual GEP but consistently underestimated mean annual ET by ~50%. Importantly, the MODIS-based models captured only 20-30% of interannual variation magnitude. These results suggest the contribution of this dryland region to variability of regional to global CO2 exchange may be up to 3-5 times larger than current estimates.

Carbon isotopic composition of forest soil respiration in the decade following bark beetle and stem girdling disturbances in the Rocky Mountains.

Bark beetle outbreaks are widespread in western North American forests, reducing primary productivity and transpiration, leading to forest mortality across large areas and altering ecosystem carbon cycling. Here the carbon isotope composition (δ(13) C) of soil respiration (δJ ) was monitored in the decade after disturbance for forests affected naturally by mountain pine beetle infestation and artificially by stem girdling. The seasonal mean δJ changed along both chronosequences. We found (a) enrichment of δJ relative to controls (<1 ‰) in near-surface soils in the first 2 years after disturbance; (b) depletion (1‰ or no change) during years 3-7; and (c) a second period of enrichment (1-2‰) in years 8-10. Results were consistent with isotopic patterns associated with the gradual death and decomposition of rhizosphere organisms, fine roots, conifer needles and woody roots and debris over the course of a decade after mortality. Finally, δJ was progressively more (13) C-depleted deeper in the soil than near the surface, while the bulk soil followed the well-established pattern of (13) C-enrichment at depth. Overall, differences in δJ between mortality classes (<1‰) and soil depths (<3‰) were smaller than variability within a class or depth over a season (up to 6‰).

Terrestrial carbon balance in a drier world: the effects of water availability in southwestern North America.

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO2 exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO2 exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site-years of annual eddy covariance measurements of net and gross CO2 exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 - 1000 mm in annual precipitation and records of 4-9 years each. In addition to evaluating spatial relationships among CO2 and water fluxes across sites, we separately quantified site-level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis-ET relationship was linear, suggesting ET was a better proxy for water available to drive CO2 exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site-level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long-term mean CO2 exchanges with climatic ET. Consequently, a hypothetical 100-mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm(-2) yr(-1). Most of the unexplained NEP variability was related to persistent, site-specific function, suggesting prioritization of research on slow-changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site-level responses to interannual weather can be extrapolated for prediction of CO2 exchanges over decadal and longer timescales relevant to societal response to climate change.

Nitrogen addition amplifies the nonlinear drought response of grassland productivity to extended growing-season droughts.

Understanding the response of grassland production and carbon exchange to intra-annual variation in precipitation and nitrogen addition is critical for sustainable grassland management and ecosystem restoration. We introduced growing-season drought treatments of different lengths (15, 30, 45 and 60 d drought) by delaying growing-season precipitation in a long-term nitrogen addition experiment in a low diversity meadow steppe in northeast China. Response variables included aboveground biomass (AGB), ecosystem net carbon exchange (NEE), and leaf net carbon assimilation rate (A). In unfertilized plots drought decreased AGB by 13.7% after a 45-d drought and 31.7% after a 60-d drought (47.6% in fertilized plots). Progressive increases in the drought response of NEE were also observed. The effects of N addition on the drought response of productivity increased as drought duration increased, and these responses were a function of changes in AGB and biomass allocation, particularly root to shoot ratio. However, no significant effects of drought occurred in fertilized or unfertilized plots in the growing season a year after the experiment, N addition did limit the recovery of AGB from severe drought during the remainder of the current growing season. Our results imply that chronic N enrichment could exacerbate the effects of growing-season drought on grassland productivity caused by altered precipitation seasonality under climate change, but that these effects do not carry over to the next growing season.