Climate‐change assessments project increasing precipitation variability through increased frequency of extreme events. However, the effects of interannual precipitation variance per se on ecosystem ...functioning have been largely understudied. Here, we report on the effects of interannual precipitation variability on the primary production of global drylands, which include deserts, steppes, shrublands, grasslands, and prairies and cover about 40% of the terrestrial earth surface. We used a global database that has 43 datasets, which are uniformly distributed in parameter space and each has at least 10 years of data. We found (a) that at the global scale, precipitation variability has a negative effect on aboveground net primary production. (b) Expected increases in interannual precipitation variability for the year 2,100 may result in a decrease of up to 12% of the global terrestrial carbon sink. (c) The effect of precipitation interannual variability on dryland productivity changes from positive to negative along a precipitation gradient. Arid sites with mean precipitation under 300 mm/year responded positively to increases in precipitation variability, whereas sites with mean precipitation over 300 mm/year responded negatively. We propose three complementary mechanisms to explain this result: (a) concave‐up and concave‐down precipitation–production relationships in arid vs. humid systems, (b) shift in the distribution of water in the soil profile, and (c) altered frequency of positive and negative legacies. Our results demonstrated that enhanced precipitation variability will have direct impacts on global drylands that can potentially affect the future terrestrial carbon sink.
Although projections of climate change anticipating increases in precipitation variability and extreme events are part of the public and scientific narrative, the effects of interannual precipitation variance per se on ecosystem functioning have been largely understudied. This global synthesis indicated that increased precipitation coefficient of variation decreased primary productivity in drylands globally. Arid ecosystems showed a positive response, while mesic ecosystems showed a negative response dominating the global response. Nonlinear productivity responses to precipitation provide mechanisms for contrasting responses across ecosystems. These findings have potential implications for the global carbon cycle and the provision of ecosystem services in drylands.
Carbon allocated underground through belowground net primary production represents the main input to soil organic carbon. This is of significant importance, because soil organic carbon is the ...third-largest carbon stock after oceanic and geological pools. However, drivers and controls of belowground productivity and the fraction of total carbon fixation allocated belowground remain uncertain. Here we estimate global belowground net primary productivity as the difference between satellite-based total net primary productivity and field observations of aboveground net primary production and assess climatic controls among biomes. On average, belowground carbon productivity is estimated as 24.7 Pg y−1, accounting for 46% of total terrestrial carbon fixation. Across biomes, belowground productivity increases with mean annual precipitation, although the rate of increase diminishes with increasing precipitation. The fraction of total net productivity allocated belowground exceeds 50% in a large fraction of terrestrial ecosystems and decreases from arid to humid ecosystems. This work adds to our understanding of the belowground carbon productivity response to climate change and provides a comprehensive global quantification of root/belowground productivity that will aid the budgeting and modeling of the global carbon cycle.
Although projections of precipitation change indicate increases in variability, most studies of impacts of climate change on ecosystems focused on effects of changes in amount of precipitation, ...overlooking precipitation variability effects, especially at the interannual scale. Here, we present results from a 6-y field experiment, where we applied sequences of wet and dry years, increasing interannual precipitation coefficient of variation while maintaining a precipitation amount constant. Increased precipitation variability significantly reduced ecosystem primary production. Dominant plantfunctional types showed opposite responses: perennial-grass productivity decreased by 81%, whereas shrub productivity increased by 67%. This pattern was explained by different nonlinear responses to precipitation. Grass productivity presented a saturating response to precipitation where dry years had a larger negative effect than the positive effects of wet years. In contrast, shrubs showed an increasing response to precipitation that resulted in an increase in average productivity with increasing precipitation variability. In addition, the effects of precipitation variation increased through time. We argue that the differential responses of grasses and shrubs to precipitation variability and the amplification of this phenomenon through time result from contrasting root distributions of grasses and shrubs and competitive interactions among plant types, confirmed by structural equation analysis. Under drought conditions, grasses reduce their abundance and their ability to absorb water that then is transferred to deep soil layers that are exclusively explored by shrubs. Our work addresses an understudied dimension of climate change that might lead to widespread shrub encroachment reducing the provisioning of ecosystem services to society.
Variability of above-ground net primary production (ANPP) of arid to sub-humid ecosystems displays a closer association with precipitation when considered across space (based on multiyear averages ...for different locations) than through time (based on year-to-year change at single locations). Here, we propose a theory of controls of ANPP based on four hypotheses about legacies of wet and dry years that explains space versus time differences in ANPP–precipitation relationships. We tested the hypotheses using 16 long-term series of ANPP. We found that legacies revealed by the association of current- versus previous-year conditions through the temporal series occur across all ecosystem types from deserts to mesic grasslands. Therefore, previous-year precipitation and ANPP control a significant fraction of current-year production. We developed unified models for the controls of ANPP through space and time. The relative importance of current-versus previous-year precipitation changes along a gradient of mean annual precipitation with the importance of current-year PPT decreasing, whereas the importance of previous-year PPT remains constant as mean annual precipitation increases. Finally, our results suggest that ANPP will respond to climate-change-driven alterations in water availability and, more importantly, that the magnitude of the response will increase with time.
Although precipitation interannual variability is projected to increase due to climate change, effects of changes in precipitation variance have received considerable less attention than effects of ...changes in the mean state of climate. Interannual precipitation variability effects on functional diversity and its consequences for ecosystem functioning are assessed here using a 6‐year rainfall manipulation experiment. Five precipitation treatments were switched annually resulting in increased levels of precipitation variability while maintaining average precipitation constant. Functional diversity showed a positive response to increased variability due to increased evenness. Dominant grasses decreased and rare plant functional types increased in abundance because grasses showed a hump‐shaped response to precipitation with a maximum around modal precipitation, whereas rare species peaked at high precipitation values. Increased functional diversity ameliorated negative effects of precipitation variability on primary production. Rare species buffered the effect of precipitation variability on the variability in total productivity because their variance decreases with increasing precipitation variance.
Climate gradients shape spatial variation in the richness and composition of plant communities. Given future predicted changes in climate means and variability, and likely regional variation in the ...magnitudes of these changes, it is important to determine how temporal variation in climate influences temporal variation in plant community structure. Here, we evaluated how species richness, turnover, and composition of grassland plant communities responded to interannual variation in precipitation by synthesizing long-term data from grasslands across the United States. We found that mean annual precipitation (MAP) was a positive predictor of species richness across sites, but a positive temporal relationship between annual precipitation and richness was only evident within two sites with low MAP. We also found higher average rates of species turnover in dry sites that in turn had a high proportion of annual species, although interannual rates of species turnover were surprisingly high across all locations. Annual species were less abundant than perennial species at nearly all sites, and our analysis showed that the probability of a species being lost or gained from one year to the next increased with decreasing species abundance. Bray-Curtis dissimilarity from one year to the next, a measure of species composition change that is influenced mainly by abundant species, was insensitive to precipitation at all sites. These results suggest that the richness and turnover patterns we observed were driven primarily by rare species, which comprise the majority of the local species pools at these grassland sites. These findings are consistent with the idea that short-lived and less abundant species are more sensitive to interannual climate variability than longer-lived and more abundant species. We conclude that, among grassland ecosystems, xeric grasslands are likely to exhibit the greatest responsiveness of community composition (richness and turnover) to predicted future increases in interannual precipitation variability. Over the long term, species composition may shift to reflect spatial patterns of mean precipitation; however, perennial-dominated systems will be buffered against rising interannual variation, while systems that have a large number of rare, annual species will show the greatest temporal variability in species composition in response to rising interannual variability in precipitation.
Understanding how biotic mechanisms confer stability in variable environments is a fundamental quest in ecology, and one that is becoming increasingly urgent with global change. Several mechanisms, ...notably a portfolio effect associated with species richness, compensatory dynamics generated by negative species covariance and selection for stable dominant species populations can increase the stability of the overall community. While the importance of these mechanisms is debated, few studies have contrasted their importance in an environmental context. We analyzed nine long-term data sets of grassland species composition to investigate how two key environmental factors, precipitation amount and variability, may directly influence community stability and how they may indirectly influence stability via biotic mechanisms. We found that the importance of stability mechanisms varied along the environmental gradient: strong negative species covariance occurred in sites characterized by high precipitation variability, whereas portfolio effects increased in sites with high mean annual precipitation. Instead of questioning whether compensatory dynamics are important in nature, our findings suggest that debate should widen to include several stability mechanisms and how these mechanisms vary in importance across environmental gradients.
Climatic changes are altering Earth's hydrological cycle, resulting in altered precipitation amounts, increased interannual variability of precipitation, and more frequent extreme precipitation ...events. These trends will likely continue into the future, having substantial impacts on net primary productivity (NPP) and associated ecosystem services such as food production and carbon sequestration. Frequently, experimental manipulations of precipitation have linked altered precipitation regimes to changes in NPP. Yet, findings have been diverse and substantial uncertainty still surrounds generalities describing patterns of ecosystem sensitivity to altered precipitation. Additionally, we do not know whether previously observed correlations between NPP and precipitation remain accurate when precipitation changes become extreme. We synthesized results from 83 case studies of experimental precipitation manipulations in grasslands worldwide. We used meta‐analytical techniques to search for generalities and asymmetries of aboveground NPP (ANPP) and belowground NPP (BNPP) responses to both the direction and magnitude of precipitation change. Sensitivity (i.e., productivity response standardized by the amount of precipitation change) of BNPP was similar under precipitation additions and reductions, but ANPP was more sensitive to precipitation additions than reductions; this was especially evident in drier ecosystems. Additionally, overall relationships between the magnitude of productivity responses and the magnitude of precipitation change were saturating in form. The saturating form of this relationship was likely driven by ANPP responses to very extreme precipitation increases, although there were limited studies imposing extreme precipitation change, and there was considerable variation among experiments. This highlights the importance of incorporating gradients of manipulations, ranging from extreme drought to extreme precipitation increases into future climate change experiments. Additionally, policy and land management decisions related to global change scenarios should consider how ANPP and BNPP responses may differ, and that ecosystem responses to extreme events might not be predicted from relationships found under moderate environmental changes.
Future changes in precipitation will strongly impact ecosystem functioning and services through changes in plant growth. Here, we synthesize 83 precipitation experiments to look at responses of above and belowground plant growth (ANPP and BNPP) across climatic gradients and levels of precipitation change extremity. Overall, we found that (1) ANPP was more responsive to precipitation increases than decreases, and this was especially evident in dry ecosystems; (2) BNPP responses were similar under precipitation increases vs. decreases; (3) under extreme wet conditions, NPP responses leveled off, creating a saturating function of NPP response vs. the magnitude of precipitation change. Based on these findings, we suggest that future research focus on BNPP and plant responses to extreme precipitation change.
Climate change is projected to cause shifts in precipitation regimes globally, leading to intensified periods of precipitation and droughts. Most studies that have explored the influence of changing ...precipitation regimes on ecosystems have focused on changes in mean annual precipitation, rather than the variance around the mean. Soil fungi are ubiquitous organisms that drive ecosystem processes, but it is unknown how they respond to long‐term increased interannual precipitation variability.
Here, we investigated the influence of long‐term increased precipitation variability and host type on soil fungal diversity and community composition in a dryland ecosystem. We collected 300 soil samples from two time points and different host type substrate types at a long‐term precipitation variability experiment at the Jornada Long Term Ecological Research site. Next, we used amplicon sequencing to characterize soil fungal communities.
Soil fungal alpha diversity and community composition were strongly affected by host type and sampling year, and increased precipitation variability caused a modest, statistically insignificant, decrease in soil fungal evenness. Furthermore, results from our structural equational model showed that the decrease in grass‐associated soil fungal richness was likely an indirect result of host decline in response to increased precipitation variability.
Synthesis. Our work demonstrates effects of increase in interannual precipitation variability on soil fungi, and that plant hosts play a key role in mediating soil fungal responses.
Our work demonstrates effects of increase in interannual precipitation variability on soil fungi, and that plant hosts play a key role in mediating soil fungal responses.
We have explored species–specific preferences for nitrate $\left({\mathrm{N}\mathrm{O}}_{3}^{-}\right)$ and ammonium $\left({\mathrm{N}\mathrm{H}}_{4}^{+}\right)$ as an alternative niche separation ...in ecosystems where nitrogen (N) is present mostly in inorganic forms. The Patagonian steppe is dominated by shrubs and grasses. Shrubs absorb water and nutrients from deep soil layers, which are poor in N, while grasses have the opposite pattern, absorbing most of their water and nutrients from the upper layers of the soil. We hypothesized that the preferences of shrub and grass for inorganic N forms are different and that the rate of potential N uptake is greater in shrubs than in grasses. To test this hypothesis, we grew individuals of six dominant species in solutions of different ${\mathrm{N}\mathrm{H}}_{4}^{+}:{\mathrm{N}\mathrm{O}}_{3}^{-}$ concentration ratios. Nitrate uptake was found to be higher in shrubs, while ammonium uptake was similar between plant functional types. The ${\mathrm{N}\mathrm{H}}_{4}^{+}:{\mathrm{N}\mathrm{O}}_{3}^{-}$ uptake ratio was significantly lower for shrubs than grasses. Shrubs, which under field conditions have deeper rooting systems than grasses, showed a higher N absorption capacity than grasses and a preference for the more mobile N form, nitrate. Grasses, which had lower N uptake rates than shrubs, preferred ammonium over nitrate. These complementary patterns between grasses and shrubs suggest a more thorough exploitation of resources by diverse ecosystems than those dominated by just one functional type. The loss of one plant functional group or a significant change in its abundance would therefore represent a reduction in resource use efficiency and ecosystem functioning.