Hotspots of biogeochemical activity linked to aridity and plant traits across global drylands.

Perennial plants create productive and biodiverse hotspots, known as fertile islands, beneath their canopies. These hotspots largely determine the structure and functioning of drylands worldwide. Despite their ubiquity, the factors controlling fertile islands under conditions of contrasting grazing by livestock, the most prevalent land use in drylands, remain virtually unknown. Here we evaluated the relative importance of grazing pressure and herbivore type, climate and plant functional traits on 24 soil physical and chemical attributes that represent proxies of key ecosystem services related to decomposition, soil fertility, and soil and water conservation. To do this, we conducted a standardized global survey of 288 plots at 88 sites in 25 countries worldwide. We show that aridity and plant traits are the major factors associated with the magnitude of plant effects on fertile islands in grazed drylands worldwide. Grazing pressure had little influence on the capacity of plants to support fertile islands. Taller and wider shrubs and grasses supported stronger island effects. Stable and functional soils tended to be linked to species-rich sites with taller plants. Together, our findings dispel the notion that grazing pressure or herbivore type are linked to the formation or intensification of fertile islands in drylands. Rather, our study suggests that changes in aridity, and processes that alter island identity and therefore plant traits, will have marked effects on how perennial plants support and maintain the functioning of drylands in a more arid and grazed world.

Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Grazing and ecosystem service delivery in global drylands.

Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure.

Cross-Site Comparisons of Dryland Ecosystem Response to Climate Change in the US Long-Term Ecological Research Network.

Long-term observations and experiments in diverse drylands reveal how ecosystems and services are responding to climate change. To develop generalities about climate change impacts at dryland sites, we compared broadscale patterns in climate and synthesized primary production responses among the eight terrestrial, nonforested sites of the United States Long-Term Ecological Research (US LTER) Network located in temperate (Southwest and Midwest) and polar (Arctic and Antarctic) regions. All sites experienced warming in recent decades, whereas drought varied regionally with multidecadal phases. Multiple years of wet or dry conditions had larger effects than single years on primary production. Droughts, floods, and wildfires altered resource availability and restructured plant communities, with greater impacts on primary production than warming alone. During severe regional droughts, air pollution from wildfire and dust events peaked. Studies at US LTER drylands over more than 40 years demonstrate reciprocal links and feedbacks among dryland ecosystems, climate-driven disturbance events, and climate change.

Precipitation versus temperature as phenology controls in drylands.

Cycles of plant growth, termed phenology, are tightly linked to environmental controls. The length of time spent growing, bounded by the start and end of season, is an important determinant of the global carbon, water, and energy balance. Much focus has been given to global warming and consequences for shifts in growing-season length in temperate regions. In conjunction with warming temperatures, altered precipitation regimes are another facet of climate change that have potentially larger consequences than temperature in dryland phenology globally. We experimentally manipulated incoming precipitation in a semiarid grassland for over a decade and recorded plant phenology at the daily scale for 7 years. We found precipitation to have a strong relationship with the timing of grass greenup and senescence but temperature had only a modest effect size on grass greenup. Pre-season drought strongly resulted in delayed grass greenup dates and shorter growing-season lengths. Spring and summer drought corresponded with earlier grass senescence, whereas higher precipitation accumulation over these seasons corresponded with delayed grass senescence. However, extremely wet conditions diluted this effect and caused a plateaued response. Deep-rooted woody shrubs showed few effects of variable precipitation or temperature on phenology and displayed consistent annual phenological timing compared with grasses. Whereas rising temperatures have already elicited phenological consequences and extended growing-season length for mid and high-latitude ecosystems, precipitation change will be the major driver of phenological change in drylands that cover 40% of the land surface with consequences for the global carbon, water, and energy balance.

Seedling responses to soil moisture amount versus pulse frequency in a successfully encroaching semi-arid shrub.

Rainfall timing, frequency, and quantity is rapidly changing in dryland regions, altering dryland plant communities. Understanding dryland plant responses to future rainfall scenarios is crucial for implementing proactive management strategies, particularly in light of land cover changes concurrent with climate change. One such change is woody plant encroachment, an increasing abundance of woody plants in areas formerly dominated by grasslands or savannas. Continued woody plant encroachment will depend, in part, on seedling capacity to establish and thrive under future climate conditions. Seedling performance is primarily impacted by soil moisture conditions governed by precipitation amount (quantity) and frequency. We hypothesized that (H1) seedling performance would be enhanced by both greater soil moisture and pulse frequency, such that seedlings with similar mean soil moisture would perform best under high pulse frequency. Alternatively, (H2) mean soil moisture would have greater influence than pulse frequency, such that a given pulse frequency would have little influence on seedling performance. The hypotheses were tested with Prosopis velutina, a shrub native to the United States that has encroached throughout its range and is invasive in other continents. Seedlings were grown in a greenhouse under two soil moisture treatments, each which was maintained by two pulse frequency treatments. Contrary to H1, mean soil moisture had greater impact than pulse frequency on seedling growth, photosynthetic gas exchange, leaf chemistry, and biomass allocation. These results indicate that P. velutina seedlings may be more responsive to rainfall amount than frequency, at least within the conditions seedlings experienced in this experimental manipulation.

Woody-plant encroachment: Precipitation, herbivory, and grass-competition interact to affect shrub recruitment.

Woody-plant encroachment is a global phenomenon that has been affecting the southwestern United States since the late 1800s. Drought, overgrazing, herbivory, and competition between grasses and shrub seedlings have been hypothesized as the main drivers of shrub establishment. However, there is limited knowledge about the interactions among these drivers. Using a rainfall manipulation system and various herbivore exclosures, we tested hypotheses about how precipitation (PPT), competition between grasses and shrub seedlings, and predation affect the germination and first-year survival of mesquite (Prosopis glandulosa), a shrub that has encroached in Southern Great Plains and Chihuahuan Desert grasslands. We found that mesquite germination and survival (1) increased with increasing PPT, then saturated at about the mean growing season PPT level, (2) that competition between grasses and shrub seedlings had no effect on either germination or survival, and (3) that herbivory by small mammals decreased seedling establishment and survival, while ant granivory showed no effect. In addition to its direct positive effect on survival, PPT had an indirect negative effect via increasing small mammal activity. Current models predict a decrease in PPT in the southwestern United States with increased frequency of extreme events. The non-linear nature of PPT effects on Mesquite recruitment suggests asymmetric responses, wherein drought has a relatively greater negative effect than the positive effect of wet years. Indirect effects of PPT, through its effects on small mammal abundance, highlight the importance of accounting for interactions between biotic and abiotic drivers of shrub encroachment. This study provides quantitative basis for developing tools that can inform effective shrub management strategies in grasslands and savannas.

Precipitation effects on nematode diversity and carbon footprint across grasslands.

Free-living nematodes are one of the most diverse metazoan taxa in terrestrial ecosystems and are critical to the global soil carbon (C) cycling through their role in organic matter decomposition. They are highly dependent on water availability for movement, feeding, and reproduction. Projected changes in precipitation across temporal and spatial scales will affect free-living nematodes and their contribution to C cycling with unforeseen consequences. We experimentally reduced and increased growing season precipitation for 2 years in 120 field plots at arid, semiarid, and mesic grasslands and assessed precipitation controls on nematode genus diversity, community structure, and C footprint. Increasing annual precipitation reduced nematode diversity and evenness over time at all sites, but the mechanism behind these temporal responses differed for dry and moist grasslands. In arid and semiarid sites, there was a loss of drought-adapted rare taxa with increasing precipitation, whereas in mesic conditions increases in the population of predaceous taxa with increasing precipitation may have caused the observed reductions in dominant colonizer taxa and yielded the negative precipitation-diversity relationship. The effects of temporal changes in precipitation on all aspects of the nematode C footprint (respiration, production, and biomass C) were all dependent on the site (significant spatial × temporal precipitation interaction) and consistent with diversity responses at mesic, but not at arid and semiarid, grasslands. These results suggest that free-living nematode biodiversity and their C footprint will respond to climate change-driven shifts in water availability and that more frequent extreme wet years may accelerate decomposition and C turnover in semiarid and arid grasslands.

Global patterns and climatic controls of belowground net carbon fixation.

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.

Ecto- and endoparasitic nematodes respond differently across sites to changes in precipitation.

Plant parasitic nematodes are among the greatest consumers of primary production in terrestrial ecosystems. Their feeding strategies can be divided into endoparasites and ectoparasites that differ substantially, not only in their damage potential to host tissue and primary production, but also in their susceptibility to environmental changes. Climate change is predicted to increase variability of precipitation in many systems, yet the effects on belowground biodiversity and associated impacts on primary productivity remain poorly understood. To examine the impact of altered precipitation on endo- and ectoparasitic soil nematodes, we conducted a 2-year precipitation manipulation study across an arid, a semiarid, and a mesic grassland. Plant parasite feeding type abundance, functional guilds, and herbivory index in response to precipitation were evaluated. Responses of endo- and ectoparasites to increased precipitation varied by grassland type. There was little response of ectoparasites to increased precipitation although their population declined at the mesic site with increased precipitation. The abundance of endoparasites remained unchanged with increasing precipitation at the arid site, increased at the semiarid, and decreased at the mesic site. The herbivory index followed closely the trends seen in the endoparasites response by stagnating at the arid site, increasing at the semiarid, and decreasing at the mesic site. Our findings suggest that altered precipitation has differing effects on plant parasite feeding strategies as well as functional guilds. This may have important implications for grassland productivity, as plant parasite pressure may exacerbate the effects of climate change on host plants.

Woody Plant Encroachment has a Larger Impact than Climate Change on Dryland Water Budgets.

Woody plant encroachment (WPE) into grasslands is a global phenomenon that is associated with land degradation via xerification, which replaces grasses with shrubs and bare soil patches. It remains uncertain how the global processes of WPE and climate change may combine to impact water availability for ecosystems. Using a process-based model constrained by watershed observations, our results suggest that both xerification and climate change augment groundwater recharge by increasing channel transmission losses at the expense of plant available water. Conversion from grasslands to shrublands without creating additional bare soil, however, reduces transmission losses. Model simulations considering both WPE and climate change are used to assess their relative roles in a late 21st century condition. Results indicate that changes in focused channel recharge are determined primarily by the WPE pathway. As a result, WPE should be given consideration when assessing the vulnerability of groundwater aquifers to climate change.

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.

Drought suppresses soil predators and promotes root herbivores in mesic, but not in xeric grasslands.

Precipitation changes among years and locations along gradients of mean annual precipitation (MAP). The way those changes interact and affect populations of soil organisms from arid to moist environments remains unknown. Temporal and spatial changes in precipitation could lead to shifts in functional composition of soil communities that are involved in key aspects of ecosystem functioning such as ecosystem primary production and carbon cycling. We experimentally reduced and increased growing-season precipitation for 2 y in field plots at arid, semiarid, and mesic grasslands to investigate temporal and spatial precipitation controls on the abundance and community functional composition of soil nematodes, a hyper-abundant and functionally diverse metazoan in terrestrial ecosystems. We found that total nematode abundance decreased with greater growing-season precipitation following increases in the abundance of predaceous nematodes that consumed and limited the abundance of nematodes lower in the trophic structure, including root feeders. The magnitude of these nematode responses to temporal changes in precipitation increased along the spatial gradient of long-term MAP, and significant effects only occurred at the mesic site. Contrary to the temporal pattern, nematode abundance increased with greater long-term MAP along the spatial gradient from arid to mesic grasslands. The projected increase in the frequency of extreme dry years in mesic grasslands will therefore weaken predation pressure belowground and increase populations of root-feeding nematodes, potentially leading to higher levels of plant infestation and plant damage that would exacerbate the negative effect of drought on ecosystem primary production and C cycling.

Changes in belowground biodiversity during ecosystem development.

Belowground organisms play critical roles in maintaining multiple ecosystem processes, including plant productivity, decomposition, and nutrient cycling. Despite their importance, however, we have a limited understanding of how and why belowground biodiversity (bacteria, fungi, protists, and invertebrates) may change as soils develop over centuries to millennia (pedogenesis). Moreover, it is unclear whether belowground biodiversity changes during pedogenesis are similar to the patterns observed for aboveground plant diversity. Here we evaluated the roles of resource availability, nutrient stoichiometry, and soil abiotic factors in driving belowground biodiversity across 16 soil chronosequences (from centuries to millennia) spanning a wide range of globally distributed ecosystem types. Changes in belowground biodiversity during pedogenesis followed two main patterns. In lower-productivity ecosystems (i.e., drier and colder), increases in belowground biodiversity tracked increases in plant cover. In more productive ecosystems (i.e., wetter and warmer), increased acidification during pedogenesis was associated with declines in belowground biodiversity. Changes in the diversity of bacteria, fungi, protists, and invertebrates with pedogenesis were strongly and positively correlated worldwide, highlighting that belowground biodiversity shares similar ecological drivers as soils and ecosystems develop. In general, temporal changes in aboveground plant diversity and belowground biodiversity were not correlated, challenging the common perception that belowground biodiversity should follow similar patterns to those of plant diversity during ecosystem development. Taken together, our findings provide evidence that ecological patterns in belowground biodiversity are predictable across major globally distributed ecosystem types and suggest that shifts in plant cover and soil acidification during ecosystem development are associated with changes in belowground biodiversity over centuries to millennia.

Effect of interannual precipitation variability on dryland productivity: A global synthesis.

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.

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.

Asynchrony among local communities stabilises ecosystem function of metacommunities.

Temporal stability of ecosystem functioning increases the predictability and reliability of ecosystem services, and understanding the drivers of stability across spatial scales is important for land management and policy decisions. We used species-level abundance data from 62 plant communities across five continents to assess mechanisms of temporal stability across spatial scales. We assessed how asynchrony (i.e. different units responding dissimilarly through time) of species and local communities stabilised metacommunity ecosystem function. Asynchrony of species increased stability of local communities, and asynchrony among local communities enhanced metacommunity stability by a wide range of magnitudes (1-315%); this range was positively correlated with the size of the metacommunity. Additionally, asynchronous responses among local communities were linked with species' populations fluctuating asynchronously across space, perhaps stemming from physical and/or competitive differences among local communities. Accordingly, we suggest spatial heterogeneity should be a major focus for maintaining the stability of ecosystem services at larger spatial scales.