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  1. Abstract Grasslands cover approximately a third of the Earth’s land surface and account for about a third of terrestrial carbon storage. Yet, we lack strong predictive models of grassland plant biomass, the primary source of carbon in grasslands. This lack of predictive ability may arise from the assumption of linear relationships between plant biomass and the environment and an underestimation of interactions of environmental variables. Using data from 116 grasslands on six continents, we show unimodal relationships between plant biomass and ecosystem characteristics, such as mean annual precipitation and soil nitrogen. Further, we found that soil nitrogen and plant diversity interacted in their relationships with plant biomass, such that plant diversity and biomass were positively related at low levels of nitrogen and negatively at elevated levels of nitrogen. Our results show that it is critical to account for the interactive and unimodal relationships between plant biomass and several environmental variables to accurately include plant biomass in global vegetation and carbon models. 
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    Free, publicly-accessible full text available December 1, 2026
  2. ABSTRACT Intermittent streams are prevalent worldwide, yet the understanding of drivers of their changing flow patterns remains incomplete. We examined hydrological changes spanning four decades (1982–2020) in Kings Creek, an intermittent grassland stream within the Konza Prairie Biological Station in Kansas, USA. We analysed streamflow data from a US Geological Survey gauge on Kings Creek and three upstream Long Term Ecological Reasearch (LTER) sub‐watersheds with annual, biennial or quadrennial burn frequencies and linked trajectories of woody encroachment to increased evapotranspiration and changes in streamflow. Riparian woody cover doubled in the annually and biannually burned sub‐watersheds and sevenfold in the quadrennially burned watersheds. We observed significant decreases (84%) in daily discharge and number of annual flow days (55%) at the downstream USGS Kings Creek gauge, with similar changes in the LTER sub‐watersheds. The changing riparian cover, propelled by the regional expansion of woody plants, contributed to decreased streamflow by amplifying actual evapotranspiration (ET). Seasonal assessments underscored the critical influence of late summer conditions (July–September), under which increases in ET were linked to rising temperatures and increased evapotranspiration by riparian cover. Our results highlight the significant hydrological impacts of woody encroachment in grasslands and emphasize the importance of long‐term ecohydrological monitoring in unravelling the interplay between climate and vegetation as controls on the hyper‐variable flow patterns in this intermittent stream. Predicting and managing hydrological impacts on the flow of intermittent grassland rivers and streams worldwide requires accounting for the effects of accelerating woody encroachment. 
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    Free, publicly-accessible full text available September 1, 2026
  3. ABSTRACT AimsThe community composition of native and alien plant species is influenced by the environment (e.g., nutrient addition and changes in temperature or precipitation). A key objective of our study is to understand how differences in the traits of alien and native species vary across diverse environmental conditions. For example, the study examines how changes in nutrient availability affect community composition and functional traits, such as specific leaf area and plant height. Additionally, it seeks to assess the vulnerability of high‐nutrient environments, such as grasslands, to alien species colonization and the potential for alien species to surpass natives in abundance. Finally, the study explores how climatic factors, including temperature and precipitation, modulate the relationship between traits and environmental conditions, shaping species success. LocationIn our study, we used data from a globally distributed experiment manipulating nutrient supplies in grasslands worldwide (NutNet). MethodsWe investigate how temporal shifts in the abundance of native and alien species are influenced by species‐specific functional traits, including specific leaf area (SLA) and leaf nutrient concentrations, as well as by environmental conditions such as climate and nutrient treatments, across 17 study sites. Mixed‐effects models were used to assess these relationships. ResultsAlien and native species increasing in their abundance did not differ in their leaf traits. We found significantly lower specific leaf area (SLA) with an increase in mean annual temperature and lower leaf Potassium with mean annual precipitation. For trait–environment relationships, when compared to native species, successful aliens exhibited an increase in leaf Phosphorus and a decrease in leaf Potassium with an increase in mean annual precipitation. Finally, aliens' SLA decreased in plots with higher mean annual temperatures. ConclusionsTherefore, studying the relationship between environment and functional traits may portray grasslands' dynamics better than focusing exclusively on traits of successful species, per se. 
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  4. Abstract ContextThe > 25,000 km2Flint Hills ecoregion in eastern Kansas and northeastern Oklahoma, USA, is an economically and ecologically important area encompassing the largest remaining tallgrass prairie ecosystem in North America. Prescribed fires are used routinely to control invasive woody species and improve forage production for the beef-cattle industry. However, burning releases harmful pollutants that, at times, contribute to air quality problems for communities across a multi-state area. ObjectivesEstablish a modeling framework for synthesizing long-term ecological data in support of Flint Hills tallgrass prairie management goals for identifying how much, where, and when rangeland burning can be conducted to maximize ecological and economic benefits while minimizing regional air quality impacts. MethodsWe used EPA’s VELMA ecohydrology model to synthesize long-term experimental data at the 35 km2Konza Prairie Biological Station (KPBS) describing the effects of climate, fire, grazing, topography, and soil moisture and nutrient dynamics on tallgrass prairie productivity and fuel loads; and to spatially extrapolate that synthesis to estimate grassland productivity and fuel loads across the nearly 1000 times larger Flint Hills ecoregion to support prescribed burning smoke trajectory modeling using the State of Kansas implementation of the U.S. Forest Service BlueSky framework. ResultsVELMA provided a performance-tested synthesis of KPBS data from field observations and experiments, thereby establishing a tool for regionally simulating the combined effects of climate, fire, grazing, topography, soil moisture, and nutrients on tallgrass prairie productivity and fuel loads. VELMA’s extrapolation of that synthesis allowed difficult-to-quantify fuel loads to be mapped across the Flint Hills to support environmental decision making, such as forecasting when, where, and how prescribed burning will have the least impact on downwind population centers. ConclusionsOur regional spatial and temporal extrapolation of VELMA’s KPBS data synthesis posits that the effects of integrated ecohydrological processes operate similarly across tallgrass prairie spatial scales. Based on multi-scale performance tests of the VELMA-BlueSky toolset, our multi-institution team is confident that it can assist stakeholders and decision makers in realistically exploring tallgrass prairie management options for balancing air quality, tallgrass prairie sustainability, and associated economic benefits for the Flint Hills ecoregion and downwind communities. 
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    Free, publicly-accessible full text available February 1, 2026
  5. Abstract In the Central Great Plains of North America, fire suppression is causing transitions from grasslands to shrublands and woodlands. This woody encroachment alters plant community composition, decreases grassland biodiversity, undermines key ecosystem services, and is difficult to reverse. How native grazers affect woody encroachment is largely unknown, especially compared to domesticated grazers. Bison were once the most widespread megafauna in North America and are typically categorized as grazers, with negative effects on grasses that indirectly benefit woody plants. However, bison can negatively impact woody plants through occasional browsing and mechanical disturbance. This study reports on a 30‐year experiment at Konza Prairie Biological Station, a mesic grassland in the Central Great Plains of North America, under fire suppression and experimental presence/absence of bison. Based on remote sensing, deciduous tree canopy cover was lower with bison (6% grazed vs. 16% ungrazed). Shrub land cover showed no difference (42% grazed vs. 41% ungrazed), while herbaceous land cover was higher with bison (51% grazed vs. 40% ungrazed). Evergreen tree canopy cover (Juniperus virginianaL.), which decreases biodiversity and increases wildfire risk, was approximately 0% with bison compared to 4% without bison. In the survival trial ofJ. virginianaseedlings, we found a 40% overwinter mortality with bison, compared to 5% mortality without bison. Compared to ungrazed areas, native plant species richness was 97% and 38% higher in bison‐grazed uplands and lowlands, respectively. Species evenness and Shannon's index were higher in the bison treatment in uplands, but not in lowlands. Bison affected community composition, resulting in higher cover of short grass species and lower tree cover. While grazers are generally assumed to favor woody plants, we found that bison had the opposite effect at low fire frequencies. We argue that the large size of bison and their behaviors account for this pattern, including trampling, horning, and occasional browsing. From a conservation perspective, bison might hamper tree expansion and increase plant diversity in tallgrass prairies and similar grasslands. 
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    Free, publicly-accessible full text available October 1, 2026
  6. ABSTRACT MotivationHere, we make available a second version of the BioTIME database, which compiles records of abundance estimates for species in sample events of ecological assemblages through time. The updated version expands version 1.0 of the database by doubling the number of studies and includes substantial additional curation to the taxonomic accuracy of the records, as well as the metadata. Moreover, we now provide an R package (BioTIMEr) to facilitate use of the database. Main Types of Variables IncludedThe database is composed of one main data table containing the abundance records and 11 metadata tables. The data are organised in a hierarchy of scales where 11,989,233 records are nested in 1,603,067 sample events, from 553,253 sampling locations, which are nested in 708 studies. A study is defined as a sampling methodology applied to an assemblage for a minimum of 2 years. Spatial Location and GrainSampling locations in BioTIME are distributed across the planet, including marine, terrestrial and freshwater realms. Spatial grain size and extent vary across studies depending on sampling methodology. We recommend gridding of sampling locations into areas of consistent size. Time Period and GrainThe earliest time series in BioTIME start in 1874, and the most recent records are from 2023. Temporal grain and duration vary across studies. We recommend doing sample‐level rarefaction to ensure consistent sampling effort through time before calculating any diversity metric. Major Taxa and Level of MeasurementThe database includes any eukaryotic taxa, with a combined total of 56,400 taxa. Software Formatcsv and. SQL. 
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    Free, publicly-accessible full text available May 1, 2026
  7. Abstract Climate change is increasing the frequency and severity of droughts globally, and grasslands are particularly vulnerable to such hydrological extremes. Drought effects at the ecosystem scale have been assessed both experimentally and through the study of naturally occurring drought, with emerging evidence that the magnitude of drought effects may vary depending on the approach used. We took advantage of a decadal study of four grasslands to directly contrast responses of aboveground net primary productivity (ANPP) to simulated vs. natural drought. The grasslands spanned a ~ threefold mean annual precipitation gradient (335–857 mm) and were all subjected to a natural 1-year drought (~ 40% reduction in precipitation from the long-term mean) and a 4 year experimental drought (~ 50% precipitation reduction). We expected that the 4 year drought would reduce ANPP more, and that post-drought recovery would be delayed, compared to the 1-year drought. We found instead that the short-term natural drought reduced ANPP more strongly than the simulated drought in all grasslands (~ 10 to ~ 50%) likely due to the co-occurrence of higher temperatures and vapor pressure deficits with reduced precipitation. Post-drought recovery was site specific and each site differed in their recovery from the natural and experimental droughts. These results align with past analyses that experiments that only manipulate soil moisture likely underestimate the magnitude of natural drought events. However, experiments can provide valuable insight into the relative sensitivity of ecosystems to reduced precipitation and soil moisture, a key aspect of drought. 
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    Free, publicly-accessible full text available July 1, 2026
  8. Abstract The size and spatial distribution of soil structural macropores impact the infiltration, percolation, and retention of soil water. Despite the assumption often made in hydrologic flux equations that these macropores are rigid, highly structured soils can respond quickly to moisture variability‐induced shrink‐swell processes altering the size distribution of these pores. In this study, we use a high‐resolution (180 m) laser imaging technique to measure the average width of interpedal, planar macropores from intact cross sections and relate it to matrix water content. We also develop an expression for unsaturated hydraulic conductivity that accounts for dynamic macropore geometries and propose a method for partitioning sensor soil water content data into matrix and macropore water contents. The model was applied to a soil in northeastern Kansas where soil monoliths had been imaged to quantify macropore properties and continuous water content data were collected at three depths. Model‐predicted macropore width showed significant sensitivity to matrix water content resulting in changes of 15%–50% of maximum width over the 15‐month period of record. Transient saturated hydraulic conductivity predicted from the model compared favorably to a previously developed model accounting for moisture‐induced changes to structural unit porosity. Following periods of low soil moisture, infiltrating meteoric water filled highly conductive macropores increasing by several orders of magnitude which subsequently decreased as water was absorbed into the matrix and macropores drained. This model offers a means by which to combine measurable morphological data with soil moisture sensors to monitor dynamic hydraulic properties of soils susceptible to shrink‐swell processes. 
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    Free, publicly-accessible full text available September 1, 2026
  9. Summary It has been 60 years since the discovery of C4photosynthesis, an event that rewrote our understanding of plant adaptation, ecosystem responses to global change, and global food security. Despite six decades of research, one aspect of C4photosynthesis that remains poorly understood is how the pathway fits into the broader context of adaptive trait spectra, which form our modern view of functional trait ecology. The C4CO2‐concentrating mechanism supports a general C4plant phenotype capable of fast growth and high resource‐use efficiencies. The fast‐efficient C4phenotype has the potential to operate at high productivity rates, while allowing for less biomass allocation to root production and nutrient acquisition, thereby providing opportunities for the evolution of novel trait covariances and the exploitation of new ecological niches. We propose the placement of the C4fast‐efficient phenotype near the acquisitive pole of the world‐wide leaf economic spectrum, but with a pathway‐specific span of trait space, wherein selection shapes both acquisitive and conservative adaptive strategies. A trait‐based perspective of C4photosynthesis will open new paths to crop improvement, global biogeochemical modeling, the management of invasive species, and the restoration of disturbed ecosystems, particularly in grasslands. 
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    Free, publicly-accessible full text available May 1, 2026
  10. Abstract Plant‐microbial‐herbivore interactions play a crucial role in the structuring and maintenance of plant communities and biodiversity, yet these relationships are complex. In grassland ecosystems, herbivores have the potential to greatly influence the survival, growth and reproduction of plants. However, few studies examine interactions of above‐ and below‐ground grazing and arbuscular mycorrhizal (AM) mycorrhizal symbiosis on plant community structure.We established experimental mesocosms containing an assemblage of eight tallgrass prairie grass and forb species in native prairie soil, maintained under mycorrhizal and nonmycorrhizal conditions, with and without native herbivorous soil nematodes, and with and without grasshopper herbivory. Using factorial analysis of variance and principal component analysis, we examined: (a) the independent and interacting effects of above‐ and below‐ground herbivores on AM symbiosis in tallgrass prairie mesocosms, (b) independent and interacting effects of above‐ and below‐ground herbivores and mycorrhizal fungi on plant community structure and (c) potential influences of mycorrhizal responsiveness of host plants on herbivory tolerance and concomitant shifts in plant community composition.Treatment effects were characterized by interactions between AM fungi and both above‐ground and below‐ground herbivores, while herbivore effects were additive. The dominance of mycorrhizal‐dependent C4grasses in the presence of AM symbiosis was increased (p < 0.0001) by grasshopper herbivory but reduced (p < 0.0001) by nematode herbivory. Cool‐season C3grasses exhibited a competitive release in the absence of AM symbiosis but this effect was largely reversed in the presence of grasshopper herbivory. Forbs showed species‐specific responses to both AM fungal inoculation and the addition of herbivores. Biomass of the grazing‐avoidant, facultatively mycotrophic forbBrickellia eupatorioidesincreased (p < 0.0001) in the absence of AM symbiosis and with grasshopper herbivory, while AM‐related increases in the above‐ground biomass of mycorrhizal‐dependent forbsRudbeckia hirtaandSalvia azureawere eradicated (p < 0.0001) by grasshopper herbivory. In contrast, nematode herbivory enhanced (p = 0.001) the contribution ofSalvia azureato total biomass.Synthesis. Our research indicates that arbuscular mycorrhizal symbiosis is the key driver of dominance of C4grasses in the tallgrass prairie, with foliar and root herbivory being two mechanisms for maintenance of plant diversity. 
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    Free, publicly-accessible full text available June 1, 2026