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  1. Summary

    Feedbacks between plants and soil microbes form a keystone to terrestrial community and ecosystem dynamics. Recent advances in dissecting the spatial and temporal dynamics of plant–soil feedbacks (PSFs) have challenged longstanding assumptions of spatially well‐mixed microbial communities and exceedingly fast microbial assembly dynamics relative to plant lifespans. Instead, PSFs emerge from interactions that are inherently mismatched in spatial and temporal scales, and explicitly considering these spatial and temporal dynamics is crucial to understanding the contribution of PSFs to foundational ecological patterns. I propose a synthetic spatiotemporal framework for future research that pairs experimental and modeling approaches grounded in mechanism to improve predictability and generalizability of PSFs.

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  2. Abstract

    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.

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  3. Abstract

    The encroachment of woody plants into grasslands is a global phenomenon with implications for biodiversity and ecosystem function. Understanding and predicting the pace of expansion and the underlying processes that control it are key challenges in the study and management of woody encroachment. Theory from spatial population biology predicts that the occurrence and speed of expansion should depend sensitively on the nature of conspecific density dependence. If fitness is maximized at the low‐density encroachment edge, then shrub expansion should be “pulled” forward. However, encroaching shrubs have been shown to exhibit positive feedbacks, whereby shrub establishment modifies the environment in ways that facilitate further shrub recruitment and survival. In this case there may be a fitness cost to shrubs at low density causing expansion to be “pushed” from behind the leading edge. We studied the spatial dynamics of creosotebush (Larrea tridentata), which has a history of encroachment into Chihuahuan Desert grasslands over the past century. We used demographic data from observational censuses and seedling transplant experiments to test the strength and direction of density dependence in shrub fitness along a gradient of shrub density at the grass–shrub ecotone. We also used seed‐drop experiments and wind data to construct a mechanistic seed‐dispersal kernel, then connected demography and dispersal data within a spatial integral projection model (SIPM) to predict the dynamics of shrub expansion. Contrary to expectations based on potential for positive feedbacks, the shrub encroachment wave is “pulled” by maximum fitness at the low‐density front. However, the predicted pace of expansion was strikingly slow (ca. 8 cm/year), and this prediction was supported by independent resurveys of the ecotone showing little to no change in the spatial extent of shrub cover over 12 years. Encroachment speed was acutely sensitive to seedling recruitment, suggesting that this population may be primed for pulses of expansion under conditions that are favorable for recruitment. Our integration of observations, experiments, and modeling reveals not only that this ecotone is effectively stalled under current conditions but also why that is so and how that may change as the environment changes.

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  4. Abstract Aim

    Roots and rhizospheres host diverse microbial communities that can influence the fitness, phenotypes, and environmental tolerances of plants. Documenting the biogeography of these microbiomes can detect the potential for a changing environment to disrupt host‐microbe interactions, particularly in cases where microbes buffer hosts against abiotic stressors. We evaluated whether root‐associated fungi had poleward declines in diversity, tested whether fungal communities in roots shifted near host plant range edges, and determined the relative importance of environmental and host predictors of root fungal community structure.


    North American plains grasslands.


    Foundation grasses –Andropogon gerardii, Bouteloua dactyloides, B. eriopoda, B. gracilis,andSchizachyrium scopariumand root fungi.


    At each of 24 sites representing three replicate 17°–latitudinal gradients, we collected roots from 12 individuals per species along five transects spaced 10 m apart (40 m × 40 m grid). We used next‐generation sequencing of ITS2, direct fungal culturing from roots, and microscopy to survey fungi associated with grass roots.


    Root‐associated fungi did not follow the poleward declines in diversity documented for many animals and plants. Instead, host plant identity had the largest influence on fungal community structure. Edaphic factors outranked climate or host plant traits as correlates of fungal community structure; however, the relative importance of environmental predictors differed among plant species. As sampling approached host species range edges, fungal composition converged in similarity among individual plants of each grass species.

    Main conclusions

    Environmental predictors of root‐associated fungi depended strongly on host plant species identity. Biogeographic patterns in fungal composition suggested a homogenizing influence of stressors at host plant range limits. Results predict that communities of non‐mycorrhizal, root‐associated fungi in the North American plains will be more sensitive to future changes in host plant ranges and edaphic factors than to the direct effects of climate.

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  5. Abstract Background

    Countries have long been making efforts by reducing greenhouse-gas emissions to mitigate climate change. In the agreements of the United Nations Framework Convention on Climate Change, involved countries have committed to reduction targets. However, carbon (C) sink and its involving processes by natural ecosystems remain difficult to quantify.


    Using a transient traceability framework, we estimated country-level land C sink and its causing components by 2050 simulated by 12 Earth System Models involved in the Coupled Model Intercomparison Project Phase 5 (CMIP5) under RCP8.5.


    The top 20 countries with highest C sink have the potential to sequester 62 Pg C in total, among which, Russia, Canada, USA, China, and Brazil sequester the most. This C sink consists of four components: production-driven change, turnover-driven change, change in instantaneous C storage potential, and interaction between production-driven change and turnover-driven change. The four components account for 49.5%, 28.1%, 14.5%, and 7.9% of the land C sink, respectively.


    The model-based estimates highlight that land C sink potentially offsets a substantial proportion of greenhouse-gas emissions, especially for countries where net primary production (NPP) likely increases substantially and inherent residence time elongates.

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  6. abstract

    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.

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  7. Abstract

    Future climates will alter the frequency and size of rain events in drylands, potentially affecting soil microbes that generate carbon feedbacks to climate, but field tests are rare. Topsoils in drylands are commonly colonized by biological soil crusts (biocrusts), photosynthesis‐based communities that provide services ranging from soil fertilization to stabilization against erosion. We quantified responses of biocrust microbial communities to 12 years of altered rainfall regimes, with 60 mm of additional rain per year delivered either as small (5 mm) weekly rains or large (20 mm) monthly rains during the summer monsoon season. Rain addition promoted microbial diversity, suppressed the dominant cyanobacterium,Microcoleus vaginatus, and enhanced nitrogen‐fixing taxa, but did not consistently increase microbial biomass. The addition of many small rain events increased microbial biomass, whereas few, large events did not. These results alter the physiological paradigm that biocrusts are most limited by the amount of rainfall and instead predict that regimes enriched in small rain events will boost cyanobacterial biocrusts and enhance their beneficial services to drylands.

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  8. Abstract Questions

    Predicting the influence of climate change on riparian plant communities improves management strategies. The sensitivity of riparian vegetation to climate and other abiotic factors depends on interactions between properties of the ecosystem, like flood regime, and plant characteristics. To explore these interactions, we addressed three questions: (a) does the composition and diversity of riparian vegetation vary with the flood regime; (b) do abiotic correlates of vegetation, including climate and groundwater, differ between sites that flood compared to locations that did not experience floods; and (c) which plant functional groups account for differential plant community sensitivity to abiotic factors between flood regimes?


    Middle Rio Grande Valley, New Mexico.


    We used long‐term observations of plant community composition, groundwater depth, precipitation and interpolated temperature from 24 sites spanning 210 km of the Rio Grande riparian cottonwood–willow forest to explore the relative importance of climate and hydrologic correlates of riparian vegetation diversity and composition.


    Riparian plant diversity was higher at sites flooding compared to non‐flooding sites. Plant diversity positively tracked shallower groundwater depth at flooding sites, but was best predicted by intra‐annual groundwater variability at non‐flooding sites. Plant community composition correlated with groundwater depth and air temperature at all sites, but at non‐flooding sites, also with intra‐annual groundwater variability and precipitation. Relationships between native plant cover and potential environmental drivers diverged strongly between the two flood regimes; non‐native plant cover had only weak relationships with most environmental predictors.


    The current flood regime of a site determined the climate and hydrologic factors that best predicted riparian plant community composition and diversity. Relationships between plant diversity or total cover and groundwater, temperature, precipitation, or groundwater variability can change in strength or direction depending on a site's flood history, highlighting the importance of flood regime to predicting the sensitivity of riparian woodlands to future environmental change.

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  9. Abstract

    Increased nutrient inputs due to anthropogenic activity are expected to increase primary productivity across terrestrial ecosystems, but changes in allocation aboveground versus belowground with nutrient addition have different implications for soil carbon (C) storage. Thus, given that roots are major contributors to soil C storage, understanding belowground net primary productivity (BNPP) and biomass responses to changes in nutrient availability is essential to predicting carbon–climate feedbacks in the context of interacting global environmental changes. To address this knowledge gap, we tested whether a decade of nitrogen (N) and phosphorus (P) fertilization consistently influenced aboveground and belowground biomass and productivity at nine grassland sites spanning a wide range of climatic and edaphic conditions in the continental United States. Fertilization effects were strong aboveground, with both N and P addition stimulating aboveground biomass at nearly all sites (by 30% and 36%, respectively, on average). P addition consistently increased root production (by 15% on average), whereas other belowground responses to fertilization were more variable, ranging from positive to negative across sites. Site‐specific responses to P were not predicted by the measured covariates. Atmospheric N deposition mediated the effect of N fertilization on root biomass and turnover. Specifically, atmospheric N deposition was positively correlated with root turnover rates, and this relationship was amplified with N addition. Nitrogen addition increased root biomass at sites with low N deposition but decreased it at sites with high N deposition. Overall, these results suggest that the effects of nutrient supply on belowground plant properties are context dependent, particularly with regard to background N supply rates, demonstrating that site conditions must be considered when predicting how grassland ecosystems will respond to increased nutrient loading from anthropogenic activity.

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  10. Abstract

    Drylands are often characterized by a pulse dynamics framework in which episodic rain events trigger brief pulses of biological activity and resource availability that regulate primary production. In the northern Chihuahuan Desert, growing season precipitation typically comes from monsoon rainstorms that stimulate soil microbial processes like decomposition, releasing inorganic nitrogen needed by plant processes. Compared to microbes, plants require greater amounts of soil moisture, typically from larger monsoon storms predicted to become less frequent and more intense in the future. Yet field‐based studies linking rainfall pulses with soil nutrient dynamics are rare. Consequently, little is known about how changes in rainfall patterns may affect plant available nitrogen in dryland soils, particularly across temporal scales. We measured daily and seasonal responses of soil inorganic nitrogen and related parameters to experimentally applied small frequent and large infrequent rain events throughout a summer growing season in a Chihuahuan Desert grassland. Contrary to long‐standing theories around resource pulse dynamics in drylands, nitrogen availability did not pulse following experimental rain events. Moreover, large infrequent events resulted in significantly less plant available nitrogen despite causing distinct pulses of increased soil moisture availability that persisted for several days. Overall, nitrogen availability increased over the growing season, especially following small frequent rain events that also stimulated some microbial ecoenzymatic activities. Our results suggest that projected changes in climate to fewer, larger rain events could significantly impact primary production in desert grasslands by decreasing plant available nitrogen when soil moisture is least limiting to plant growth.

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