Seasonal,not annual precipitation drives community productivity across ecosystems

Understanding drivers of aboveground net primary production (ANPP) has long been a goal of ecology. Decades of investigation have shown total annual precipitation to be an important determinant of ANPP within and across ecosystems. Recently a few studies at individual sites have shown precipitation during specific seasons of the year can more effectively predict ANPP. Here we determined whether seasonal or total precipitation better predicted ANPP across a range of terrestrial ecosystems, from deserts to forests, using long‐term data from 36 plant communities. We also determined whether ANPP responses were dependent on ecosystem type or plant functional group. We found that seasonal precipitation generally explained ANPP better than total precipitation. Precipitation in multiple parts of the growing season often correlated with ANPP, but rarely interacted with each other. Surprisingly, the amount of variation explained by seasonal precipitation was not correlated with ecosystem type or plant functional group. Overall, examining seasonal precipitation can significantly improve ANPP predictions across a broad range of ecosystems and plant types, with implications for understanding current and future ANPP variation. Further work examining precipitation timing relative to species phenology may further improve our ability to predict ANPP, especially in response to climate change.     

Press–pulse interactions: effects of warming,N deposition,altered winter precipitation,and fire on desert grassland community structure and dynamics

Global environmental change is altering temperature, precipitation patterns, resource availability, and disturbance regimes. Theory predicts that ecological presses will interact with pulse events to alter ecosystem structure and function. In 2006, we established a long‐term, multifactor global change experiment to determine the interactive effects of nighttime warming, increased atmospheric nitrogen (N) deposition, and increased winter precipitation on plant community structure and aboveground net primary production (ANPP) in a northern Chihuahuan Desert grassland. In 2009, a lightning‐caused wildfire burned through the experiment. Here, we report on the interactive effects of these global change drivers on pre‐ and postfire grassland community structure and ANPP. Our nighttime warming treatment increased winter nighttime air temperatures by an average of 1.1 °C and summer nighttime air temperature by 1.5 °C. Soil N availability was 2.5 times higher in fertilized compared with control plots. Average soil volumetric water content (VWC) in winter was slightly but significantly higher (13.0% vs. 11.0%) in plots receiving added winter rain relative to controls, and VWC was slightly higher in warmed (14.5%) compared with control (13.5%) plots during the growing season even though surface soil temperatures were significantly higher in warmed plots. Despite these significant treatment effects, ANPP and plant community structure were highly resistant to these global change drivers prior to the fire. Burning reduced the cover of the dominant grasses by more than 75%. Following the fire, forb species richness and biomass increased significantly, particularly in warmed, fertilized plots that received additional winter precipitation. Thus, although unburned grassland showed little initial response to multiple ecological presses, our results demonstrate how a single pulse disturbance can interact with chronic alterations in resource availability to increase ecosystem sensitivity to multiple drivers of global environmental change.     

Nitrogen limitation in dryland ecosystems: Responses to geographical and temporal variation in precipitation

We investigated the relationship between plant nitrogen limitation and water availability in dryland ecosystems. We tested the hypothesis that at lower levels of annual precipitation, aboveground net primary productivity (ANPP) is limited primarily by water whereas at higher levels of precipitation, it is limited primarily by nitrogen. Using a literature survey of fertilization experiments in arid, semi-arid, and subhumid ecosystems, we investigated the response of ANPP to nitrogen addition as a function of variation in precipitation across geographic gradients, as well as across year-to-year variation in precipitation within sites. We used four different indices to assess the degree of N limitation: (1) Absolute Increase of plant production in response to fertilization (the slope of ANPP vs. amount of added N at different levels of annual precipitation); (2) Relative Response (the percent increase in fertilized over control ANPP at different levels of N addition); (3) Fertilizer Use Efficiency (FUE, the absolute gain in productivity per amount of fertilizer N), and (4) Maximum Response (the greatest absolute increase in ANPP at saturating levels of N addition). Relative Response to fertilization did not significantly increase with increasing precipitation either across the geographic gradient or across year-to-year variation within sites. Nor did the Maximum Response to fertilization increase with increasing precipitation across the geographic gradient. On the other hand, there was a significant increase in the Absolute Increase and FUE indices with both geographical and temporal variation in precipitation. Together, these results indicate that there is not necessarily a shift of primary limitation from water to N across the geographic water availability gradient. Instead, our results support the hypothesis of co-limitation. The apparently contradictory results from the four indices of N limitation can best be explained by an integration of plant ecophysiological, community, and ecosystem mechanisms whereby plants are co-limited by multiple resources, species shifts occur in response to changing resource levels, and nitrogen and water availability are tightly linked through biogeochemical feedbacks.     

A test of two mechanisms proposed to optimize grassland aboveground primary productivity in response to grazing

Aims Mesic grasslands have a long evolutionary history of grazing by large herbivores and as a consequence, grassland species have numerous adaptations allowing them to respond favourably to grazing. Although empirical evidence has been equivocal, theory predicts that such adaptations combined with alterations in resources can lead to grazing-induced overcompensation in aboveground net primary production (ANPP; grazed ANPP> ungrazed ANPP) under certain conditions. We tested two specific predictions from theory. First, overcompensation is more likely to occur in annually burned grasslands because limiting nutrients that would be lost with frequent fires are recycled through grazers and stimulate ANPP. Second, overcompensation of biomass lost to grazers is more likely to occur in unburned sites where grazing has the greatest effect on increasing light availability through alterations in canopy structure.Methods We tested these nutrient versus light-based predictions in grazed grasslands that had been annually burned or protected from fire for>20 years. We assessed responses in ANPP to grazing by large ungulates using both permanent and moveable grazing exclosures (252 exclosures from which biomass was harvested from 3192 quadrats) in a 2-year study. Study sites were located at the Konza Prairie Biological Station (KPBS) in North America and at Kruger National Park (KNP) in South Africa. At KPBS, sites were grazed by North American bison whereas in KNP sites were grazed either by a diverse suite of herbivores (e.g. blue wildebeest, Burchell's zebra, African buffalo) or by a single large ungulate (African buffalo).Important findings We found no evidence for overcompensation in either burned or unburned sites, regardless of grazer type. Thus, there was no support for either mechanism leading to overcompensation. Instead, complete compensation of total biomass lost to grazers was the most common response characterizing grazing–ANPP relationships with, in some cases, undercompensation of grass ANPP being offset by increased ANPP of forbs likely due to competitive release. The capability of these very different grass-dominated systems to maintain ANPP while being grazed has important implications for energy flow, ecosystem function and the trophic dynamics of grasslands.     

Shifts in the dynamics of productivity signal ecosystem state transitions at the biome‐scale

Understanding ecosystem dynamics and predicting directional changes in ecosystem in response to global changes are ongoing challenges in ecology. Here we present a framework that links productivity dynamics and ecosystem state transitions based on a spatially continuous dataset of aboveground net primary productivity (ANPP) from the temperate grassland of China. Across a regional precipitation gradient, we quantified spatial patterns in ANPP dynamics (variability, asymmetry and sensitivity to rainfall) and related these to transitions from desert to semi‐arid to mesic steppe. We show that these three indices of ANPP dynamics displayed distinct spatial patterns, with peaks signalling transitions between grassland types. Thus, monitoring shifts in ANPP dynamics has the potential for predicting ecosystem state transitions in the future. Current ecosystem models fail to capture these dynamics, highlighting the need to incorporate more nuanced ecological controls of productivity in models to forecast future ecosystem shifts.     

Regional grassland productivity responses to precipitation during multiyear above‐ and below‐average rainfall periods

There is considerable uncertainty in the magnitude and direction of changes in precipitation associated with climate change, and ecosystem responses are also uncertain. Multiyear periods of above‐ and below‐average rainfall may foretell consequences of changes in rainfall regime. We compiled long‐term aboveground net primary productivity (ANPP) and precipitation (PPT) data for eight North American grasslands, and quantified relationships between ANPP and PPT at each site, and in 1–3 year periods of above‐ and below‐average rainfall for mesic, semiarid cool, and semiarid warm grassland types. Our objective was to improve understanding of ANPP dynamics associated with changing climatic conditions by contrasting PPT–ANPP relationships in above‐ and below‐average PPT years to those that occurred during sequences of multiple above‐ and below‐average years. We found differences in PPT–ANPP relationships in above‐ and below‐average years compared to long‐term site averages, and variation in ANPP not explained by PPT totals that likely are attributed to legacy effects. The correlation between ANPP and current‐ and prior‐year conditions changed from year to year throughout multiyear periods, with some legacy effects declining, and new responses emerging. Thus, ANPP in a given year was influenced by sequences of conditions that varied across grassland types and climates. Most importantly, the influence of prior‐year ANPP often increased with the length of multiyear periods, whereas the influence of the amount of current‐year PPT declined. Although the mechanisms by which a directional change in the frequency of above‐ and below‐average years imposes a persistent change in grassland ANPP require further investigation, our results emphasize the importance of legacy effects on productivity for sequences of above‐ vs. below‐average years, and illustrate the utility of long‐term data to examine these patterns.     

Differential roles of species richness versus species asynchrony in regulating community stability along a precipitation gradient

Plant community may provide products and services to humans. However, patterns and drivers of community stability along a precipitation gradient remain unclear. A regional‐scale transect survey was conducted over a 3‐year period from 2013 to 2015, along a precipitation gradient from 275 to 555 mm and spanning 440 km in length from west to east in a temperate semiarid grassland of northern China, a central part of the Eurasian steppe. Our study provided regional‐scale evidence that the community stability increased with increasing precipitation in the semiarid ecosystem. The patterns of community stability along a precipitation gradient were ascribed to community composition and community dynamics, such as species richness and species asynchrony, rather than the abiotic effect of precipitation. Species richness regulated the temporal mean (μ) of aboveground net primary productivity (ANPP), while species asynchrony regulated the temporal standard deviation (σ) of ANPP, which in turn contributed to community stability. Our findings highlight the crucial role of community composition and community dynamics in regulating community stability under climate change.     

Aboveground net primary production dynamics in a northern Chihuahuan Desert ecosystem

Aboveground net primary production (ANPP) dynamics are a key element in the understanding of ecosystem processes. For semiarid environments, the pulse-reserve framework links ANPP to variable and unpredictable precipitation events contingent on surficial hydrology, soil moisture dynamics, biodiversity structure, trophic dynamics, and landscape context. Consequently, ANPP may be decoupled periodically from processes such as decomposition and may be subjected to complex feedbacks and thresholds at broader scales. As currently formulated, the pulse-reserve framework may not encompass the breadth of ANPP response to seasonal patterns of precipitation and heat inputs. Accordingly, we examined a 6-year (1999–2004), seasonal record of ANPP with respect to precipitation, soil moisture dynamics, and functional groups in a black grama (Bouteloua eriopoda) grassland and a creosotebush (Larrea tridentata) shrubland in the northern Chihuahuan Desert. Annual ANPP was similar in the grassland (51.1 g/m²) and shrubland (59.2 g/m²) and positively correlated with annual precipitation. ANPP differed among communities with respect to life forms and functional groups and responses to abiotic drivers. In keeping with the pulse-reserve model, ANPP in black grama grassland was dominated by warm-season C₄ grasses and subshrubs that responded to large, transient summer storms and associated soil moisture in the upper 30 cm. In contrast, ANPP in creosotebush shrubland occasionally responded to summer moisture, but the predominant pattern was slower, non-pulsed growth of cool-season C₃ shrubs during spring, in response to winter soil moisture accumulation and the breaking of cold dormancy. Overall, production in this Chihuahuan Desert ecosystem reflected a mix of warm-temperate arid land pulse dynamics during the summer monsoon and non-pulsed dynamics in spring driven by winter soil moisture accumulation similar to that of cool-temperate regions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Global‐change drivers of ecosystem functioning modulated by natural variability and saturating responses

Humans are altering global environment at an unprecedented rate through changes in biodiversity, climate, nitrogen cycle, and land use. To address their effects on ecosystem functioning, experiments most frequently explore one driver at a time and control as many confounding factors as possible. Yet, which driver exerts the largest influence on ecosystem functioning and whether their relative importance changes among systems remain unclear. We analyzed experiments in the Patagonian steppe that evaluated the aboveground net primary production (ANPP) response to manipulated gradients of species richness, precipitation, temperature, nitrogen fertilization (N), and grazing intensity. We compared the effect on ANPP relative to ambient conditions considering intensity and direction of manipulations for each driver. The ranking of responses to drivers with comparable manipulation intensity was as follows: biodiversity>grazing>precipitation>N. For a similar intensity of manipulation, the effect of biodiversity loss was 4.0, 3.6, and 1.5, times larger than N deposition, decreased precipitation, and increased grazing intensity. We interpreted our results considering two hypotheses. First, the response of ANPP to changes in precipitation and biodiversity is saturating, so we expected larger effects when the driver was reduced, relative to ambient conditions, than when it was increased. Experimental manipulations that reduced ambient levels had larger effects than those that increased them. Second, the sensitivity of ANPP to each driver is inversely related to the natural variability of the driver. In Patagonia, the ranking of natural variability of drivers is as follows: precipitation>grazing>temperature>biodiversity>N. So, in general, the ecosystem was most sensitive to drivers that varied the least. Comparable results from Cedar Creek (MN) support both hypotheses and suggest that sensitivity to drivers varies among ecosystem types. Given the importance of understanding ecosystem sensitivity to predict global‐change impacts, it is necessary to design new experiments located in regions with contrasting natural variability and that include the full range of drivers.     

Understanding the long‐term spatial dynamics of a semiarid grass‐shrub steppe through inverse parameterization for simulation models

Desertification threatens 70% of all dry lands worldwide by diminishing the provision of economic and ecosystem services. However, since long‐term vegetation dynamics of semiarid ecosystems are difficult to study, the opportunities to evaluate desertification and degradation properly are limited. In this study, we tailored, calibrated and tested a spatially‐explicit simulation model (DINVEG) to describe the long‐term dynamics of dominant grass and shrub species in the semiarid Patagonian steppe. We used inverse techniques to identify parameterizations that yield model outputs in agreement with detailed field data, and we performed sensitivity analyses to reveal the main drivers of long‐term vegetation dynamics. Whereas many parameterizations (10–45%) matched single field observations (e.g. grass and shrub cover, species‐specific density, aboveground net primary production [ANPP]), only a few parameterizations (0.05%) yielded simultaneous match of all field observations. Sensitivity analysis pointed to demographic constraints for shrubs and grasses in the emergence and recruitment phase, respectively, which contributed to balanced shrub‐grass abundances in the long run. Vegetation dynamics of simulations that matched all field observations were characterized by a stochastic equilibrium. The soil water content in the top layer (0–10 cm) during the emergence period was the strongest predictor of shrub densities and population growth rates and of growth rates of grasses. Grasses controlled the shrub demography because of the resource overlap of grasses with juvenile shrubs (i.e. water content in the top layer). In agreement with field observations, ecosystem function buffered the strong variability in precipitation (a simulated CV in ANPP of 16% vs CV in precipitation of 33%). Our results show that seedling emergence and recruitment are critical processes for long‐term vegetation dynamics in this steppe. The methods presented here could be widely applied when data for direct parameterization of individual‐based models are lacking, but data corresponding to model outputs are available. Our modeling methodology can reduce the need for long‐term data sets when answering questions regarding community dynamics.     

As above,not so below: Long-term dynamics of net primary production across a dryland transition zone

Drylands are key contributors to interannual variation in the terrestrial carbon sink, which has been attributed primarily to broad-scale climatic anomalies that disproportionately affect net primary production (NPP) in these ecosystems. Current knowledge around the patterns and controls of NPP is based largely on measurements of aboveground net primary production (ANPP), particularly in the context of altered precipitation regimes. Limited evidence suggests belowground net primary production (BNPP), a major input to the terrestrial carbon pool, may respond differently than ANPP to precipitation, as well as other drivers of environmental change, such as nitrogen deposition and fire. Yet long-term measurements of BNPP are rare, contributing to uncertainty in carbon cycle assessments. Here, we used 16 years of annual NPP measurements to investigate responses of ANPP and BNPP to several environmental change drivers across a grassland–shrubland transition zone in the northern Chihuahuan Desert. ANPP was positively correlated with annual precipitation across this landscape; however, this relationship was weaker within sites. BNPP, on the other hand, was weakly correlated with precipitation only in Chihuahuan Desert shrubland. Although NPP generally exhibited similar trends among sites, temporal correlations between ANPP and BNPP within sites were weak. We found chronic nitrogen enrichment stimulated ANPP, whereas a one-time prescribed burn reduced ANPP for nearly a decade. Surprisingly, BNPP was largely unaffected by these factors. Together, our results suggest that BNPP is driven by a different set of controls than ANPP. Furthermore, our findings imply belowground production cannot be inferred from aboveground measurements in dryland ecosystems. Improving understanding around the patterns and controls of dryland NPP at interannual to decadal scales is fundamentally important because of their measurable impact on the global carbon cycle. This study underscores the need for more long-term measurements of BNPP to improve assessments of the terrestrial carbon sink, particularly in the context of ongoing environmental change.     

Controls of Aboveground Net Primary Production in Mesic Savanna Grasslands: An Inter-Hemispheric Comparison

Patterns and controls of annual aboveground net primary productivity (ANPP) are fundamental metrics of ecosystem functioning. It is generally assumed, but rarely tested, that determinants of ANPP in one region within a biome will operate similarly throughout that biome, as long as physiognomy and climate are broadly consistent. We tested this assumption by quantifying ANPP responses to fire, grazing history, and nitrogen (N) addition in North American (NA) and South African (SA) savanna grasslands. We found that total ANPP responded in generally consistent ways to fire, grazing history, and N addition on both continents. Annual fire in both NA and SA consistently stimulated total ANPP (28–100%) relative to unburned treatments at sites with deep soils, and had no effect on ANPP in sites with shallow soils. Fire did not affect total ANPP in sites with a recent history of grazing, regardless of whether a single or a diverse suite of large herbivores was present. N addition interacted strongly and consistently with fire regime in both NA and SA. In annually burned sites that were not grazed, total ANPP was stimulated by N addition (29–39%), but there was no effect of N fertilization in the absence of fire. In contrast, responses in forb ANPP to fire and grazing were somewhat divergent across this biome. Annual fire in NA reduced forb ANPP, whereas grazing increased forb ANPP, but neither response was evident in SA. Thus, despite a consistent response in total ANPP, divergent responses in forb ANPP suggest that other aspects of community structure and ecosystem functioning differ in important ways between these mesic savanna grasslands.     

Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation

Global mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta‐analysis to synthesize ecosystem‐level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single‐factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.     

Drivers of Variation in Aboveground Net Primary Productivity and Plant Community Composition Differ Across a Broad Precipitation Gradient

Aboveground net primary production (ANPP) is a key integrator of C uptake and energy flow in many terrestrial ecosystems. As such, ecologists have long sought to understand the factors driving variation in this important ecosystem process. Although total annual precipitation has been shown to be a strong predictor of ANPP in grasslands across broad spatial scales, it is often a poor predictor at local scales. Here we examine the amount of variation in ANPP that can be explained by total annual precipitation versus precipitation during specific periods of the year (precipitation periods) and nutrient availability at three sites representing the major grassland types (shortgrass steppe, mixed-grass prairie, and tallgrass prairie) spanning the broad precipitation gradient of the U.S. Central Great Plains. Using observational data, we found that precipitation periods and nutrient availability were much stronger predictors of site-level ANPP than total annual precipitation. However, the specific nutrients and precipitation periods that best predicted ANPP differed among the three sites. These effects were mirrored experimentally at the shortgrass and tallgrass sites, with precipitation and nutrient availability co-limiting ANPP, but not at the mixed-grass site, where nutrient availability determined ANPP exclusive of precipitation effects. Dominant grasses drove the ANPP response to increased nutrient availability at all three sites. However, the relative responses of rare grasses and forbs were greater than those of the dominant grasses to experimental nutrient additions, thus potentially driving species turnover with chronic nutrient additions. This improved understanding of the factors driving variation in ANPP within ecosystems spanning the broad precipitation gradient of the Great Plains will aid predictions of alterations in ANPP under future global change scenarios.     

Successional change in species composition alters climate sensitivity of grassland productivity

Succession theory predicts altered sensitivity of ecosystem functions to disturbance (i.e., climate change) due to the temporal shift in plant community composition. However, empirical evidence in global change experiments is lacking to support this prediction. Here, we present findings from an 8‐year long‐term global change experiment with warming and altered precipitation manipulation (double and halved amount). First, we observed a temporal shift in species composition over 8 years, resulting in a transition from an annual C₃‐dominant plant community to a perennial C₄‐dominant plant community. This successional transition was independent of any experimental treatments. During the successional transition, the response of aboveground net primary productivity (ANPP) to precipitation addition magnified from neutral to +45.3%, while the response to halved precipitation attenuated substantially from ?17.6% to neutral. However, warming did not affect ANPP in either state. The findings further reveal that the time‐dependent climate sensitivity may be regulated by successional change in species composition, highlighting the importance of vegetation dynamics in regulating the response of ecosystem productivity to precipitation change.     

Environmental factors covary with plant diversity–productivity relationships among Chinese grassland sites

Aim Our objective was to document the general relationship between plant species richness (SR) and above‐ground net primary productivity (ANPP) at different spatial scales and the environmental influence on this relationship. Location Temperate and alpine grasslands of China. Methods We investigated SR and ANPP at 321 field sites (1355 plots) across the widely distributed temperate and alpine grasslands of China. Ordinary least squares (OLS) regressions were used to test SR–ANPP relationships among site means. Plot‐level data of SR and ANPP were analysed with general linear models (GLMs) and the correlation between SR and ANPP was decomposed into covariance components to test the influence of climatic variables, region, vegetation type and remaining variation among sites on SR, ANPP and their relationship. Results We found positive linear relationships between SR and ANPP among sites in both the alpine and temperate grassland regions and in different grassland vegetation types of these biomes. Environmental gradients such as growing‐season precipitation affected both SR and ANPP in parallel. However, after removing the among‐site environmental variation, residual SR and ANPP were no longer correlated at the pooled within‐site level. Main conclusions The positive SR–ANPP relationship across large‐scale environmental gradients among sites was most likely the result of climatic variables influencing SR and ANPP in parallel. Our results suggest that in China's natural grasslands there is no direct relationship between SR and ANPP, presumably because the pool of available species for local community assembly is large, in contrast to experiments where species pools are artificially reduced.     

Inter‐annual variation in primary production of a semi‐arid grassland related to previous‐year production

Mean annual precipitation accounts for a large proportion of the variation in mean above‐ground net primary production (ANPP) of grasslands worldwide. However, the inter‐annual variation in production in any grassland site is only loosely correlated with precipitation. The longest record of variation in production and precipitation for a site corresponds to a shortgrass steppe in Colorado, USA. A previous study of this record showed that current‐year precipitation accounted for 39% of the inter‐annual variation in ANPP. In this note, we show that ca. one third of the unexplained variation is related to previous‐year ANPP: ANPP per mm of precipitation was higher in years preceded by wet, more productive years than in years preceded by average years; similarly, ANPP per mm of precipitation was lower in years preceded by dry, less productive years than in years preceded by average years. Since previous‐year ANPP was, in turn, associated with precipitation of a year before, current‐year ANPP was also explained by precipitation of two previous years. Our finding not only increases our predictive ability, but it also changes our understanding of how ANPP responds to fluctuations in precipitation. If ANPP is thought to vary according to current‐year precipitation only, it will simply track annual precipitation in time. According to this new result, however, ANPP fluctuations are buffered if wet, more productive years alternate with dry, less productive years, and they are amplified if wet or dry sequences of several years take place.     

Carbon allocation in boreal black spruce forests across regions varying in soil temperature and precipitation

A common hypothesis for northern ecosystems is that low soil temperatures inhibit plant productivity. To address this hypothesis, we reviewed how separate components of ecosystem carbon (C) cycling varied along a soil temperature gradient for nine well-drained, relatively productive boreal black spruce ( Picea mariana Mill. [B.S.P.]) forests in Alaska, USA, and Saskatchewan and Manitoba, Canada. Annual soil temperature [expressed as soil summed degree days (SDD)] was positively correlated with aboveground net primary productivity (ANPP), while negatively correlated with total belowground carbon flux (TBCF). The partitioning of C to ANPP at the expense of root processes represented a nearly 1 : 1 tradeoff across the soil temperature gradient, which implied that the amount of C cycling through these black spruce ecosystems was relatively insensitive to variation in SDD. Moreover, the rate at which C accumulated in the ecosystem since the last stand replacing fire was unrelated to SDD, but SDD was positively correlated to the ratio of spruce-biomass : forest-floor-mass. Thus, plant partitioning of C and the distribution of ecosystem C were apparently affected by soil temperature, although across regions, precipitation co-varied with soil temperature. These two factors likely correlated with one another because of precipitation's influence on soil heat balance, suggesting that a soil temperature–precipitation interaction could be responsible for the shifts in C allocation. Nonetheless, our results highlight that for this boreal ecosystem, ANPP and TBCF can be negatively correlated. In tropical and temperate forests, TBCF and ANPP have been reported as positively correlated, and our results may reflect the unique interactions between soil temperature, forest floor accumulation, rooting depth, and nutrient availability that characterize the black spruce forest type.     

Sensitivity of mean annual primary production to precipitation

In many terrestrial ecosystems, variation in aboveground net primary production (ANPP) is positively correlated with variation in interannual precipitation. Global climate change will alter both the mean and the variance of annual precipitation, but the relative impact of these changes in precipitation on mean ANPP remains uncertain. At any given site, the slope of the precipitation‐ANPP relationship determines the sensitivity of mean ANPP to changes in mean precipitation, whereas the curvature of the precipitation‐ANPP relationship determines the sensitivity of ANPP to changes in precipitation variability. We used 58 existing long‐term data sets to characterize precipitation‐ANPP relationships in terrestrial ecosystems and to quantify the sensitivity of mean ANPP to the mean and variance of annual precipitation. We found that most study sites have a nonlinear, saturating relationship between precipitation and ANPP, but these nonlinearities were not strong. As a result of these weak nonlinearities, ANPP was nearly 40 times more sensitive to precipitation mean than variance. A 1% increase in mean precipitation caused a ?0.2% to 1.8% change in mean ANPP, with a 0.64% increase on average. Sensitivities to precipitation mean peaked at sites with a mean annual precipitation near 500 mm. Changes in species composition and increased intra‐annual precipitation variability could lead to larger ANPP responses to altered precipitation regimes than predicted by our analysis.     

Precipitation amount and event size interact to reduce ecosystem functioning during dry years in a mesic grassland

Ongoing intensification of the hydrological cycle is altering rainfall regimes by increasing the frequency of extreme wet and dry years and the size of individual rainfall events. Despite long‐standing recognition of the importance of precipitation amount and variability for most terrestrial ecosystem processes, we lack understanding of their interactive effects on ecosystem functioning. We quantified this interaction in native grassland by experimentally eliminating temporal variability in growing season rainfall over a wide range of precipitation amounts, from extreme wet to dry conditions. We contrasted the rain use efficiency (RUE) of above‐ground net primary productivity (ANPP) under conditions of experimentally reduced versus naturally high rainfall variability using a 32‐year precipitation–ANPP dataset from the same site as our experiment. We found that increased growing season rainfall variability can reduce RUE and thus ecosystem functioning by as much as 42% during dry years, but that such impacts weaken as years become wetter. During low precipitation years, RUE is lowest when rainfall event sizes are relatively large, and when a larger proportion of total rainfall is derived from large events. Thus, a shift towards precipitation regimes dominated by fewer but larger rainfall events, already documented over much of the globe, can be expected to reduce the functioning of mesic ecosystems primarily during drought, when ecosystem processes are already compromised by low water availability.