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  1. Abstract Background and aims

    A synergistic response of aboveground plant biomass production to combined nitrogen (N) and phosphorus (P) addition has been observed in many ecosystems, but the underlying mechanisms and their relative importance are not well known. We aimed at evaluating several mechanisms that could potentially cause the synergistic growth response, such as changes in plant biomass allocation, increased N and P uptake by plants, and enhanced ecosystem nutrient retention.


    We studied five grasslands located in Europe and the USA that are subjected to an element addition experiment composed of four treatments: control (no element addition), N addition, P addition, combined NP addition.


    Combined NP addition increased the total plant N stocks by 1.47 times compared to the N treatment, while total plant P stocks were 1.62 times higher in NP than in single P addition. Further, higher N uptake by plants in response to combined NP addition was associated with reduced N losses from the soil (evaluated based on soil δ15N) compared to N addition alone, indicating a higher ecosystem N retention. In contrast, the synergistic growth response was not associated with significant changes in plant resource allocation.


    Our results demonstrate that the commonly observed synergistic effect of NP addition on aboveground biomass production in grasslands is caused by enhanced N uptake compared to single N addition, and increased P uptake compared to single P addition, which is associated with a higher N and P retention in the ecosystem.

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  2. Abstract

    We review results from field experiments that simulate drought, an ecologically impactful global change threat that is predicted to increase in magnitude, extent, duration and frequency. Our goal is to address, from primarily an ecosystem perspective, the questions ‘What have we learned from drought experiments?’ and ‘Where do we go from here?’.

    Drought experiments are among the most numerous climate change manipulations and have been deployed across a wide range of biomes, although most are conducted in short‐statured, water‐limited ecosystems. Collectively, these experiments have enabled ecologists to quantify the negative responses to drought that occur for most aspects of ecosystem structure and function. Multiple meta‐analyses of responses have also enabled comparisons of relative effect sizes of drought across hundreds of sites, particularly for carbon cycle metrics. Overall, drought experiments have provided strong evidence that ecosystem sensitivity to drought increases with aridity, but that plant traits associated with aridity are not necessarily predictive of drought resistance. There is also intriguing evidence that as drought magnitude or duration increases to extreme levels, plant strategies may shift from drought tolerance to drought escape/avoidance.

    We highlight three areas where more drought experiments are needed to advance our understanding. First, because drought is intensifying in multiple ways, experiments are needed that address alterations in drought magnitude versus duration, timing and/or frequency (individually and interactively). Second, drivers of drought may be shifting—from precipitation deficits to rising atmospheric demand for water—and disentangling how ecosystems respond to changes in hydrological ‘supply versus demand’ is critical for understanding drought impacts in the future. Finally, more attention should be focussed on post‐drought recovery periods since legacies of drought can affect ecosystem functioning much longer than the drought itself.

    We conclude with a call for a fundamental shift in the focus of drought experiments from those designed primarily as ‘response experiments’, quantifying the magnitude of change in ecosystem structure and function, to more ‘mechanistic experiments’—those that explicitly manipulate ecological processes or attributes thought to underpin drought responses.

    Read the freePlain Language Summaryfor this article on the Journal blog.

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  3. Hui, Dafeng (Ed.)

    While the relationship between genetic diversity and plant productivity has been established for many species, it is unclear whether environmental conditions and biotic associations alter the nature of the relationship. To address this, we investigated the interactive effects of genotypic diversity, drought and mycorrhizal association on plant productivity and plant traits. Our mesocosm study was set up at the Konza Prairie Biological Research Station, located in the south of Manhattan, Kansas. Andropogon gerardii, the focal species for our study, was planted in two levels of genotypic richness treatment: monoculture or three-genotype polyculture. A rainout shelter was constructed over half of the experimental area to impose a drought and Thiophanate-methyl fungicide was used to suppress arbuscular mycorrhizal fungi in selected pots within each genotypic richness and drought treatment. Genotypic richness and mycorrhizal association did not affect above-ground biomass of A. gerardii. Drought differentially affected the above-ground biomass, the number of flowers and bolts of A. gerardii genotypes, and the biomass and the functional traits also differed for monoculture versus polyculture. Our results suggest that drought and genotypic richness can have variable outcomes for different genotypes of a plant species.

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  4. Summary

    Evolutionary history plays a key role driving patterns of trait variation across plant species. For scaling and modeling purposes, grass species are typically organized into C3vs C4plant functional types (PFTs). Plant functional type groupings may obscure important functional differences among species. Rather, grouping grasses by evolutionary lineage may better represent grass functional diversity.

    We measured 11 structural and physiological traitsin situfrom 75 grass species within the North American tallgrass prairie. We tested whether traits differed significantly among photosynthetic pathways or lineages (tribe) in annual and perennial grass species.

    Critically, we found evidence that grass traits varied among lineages, including independent origins of C4photosynthesis. Using a rigorous model selection approach, tribe was included in the top models for five of nine traits for perennial species. Tribes were separable in a multivariate and phylogenetically controlled analysis of traits, owing to coordination of important structural and ecophysiological characteristics.

    Our findings suggest grouping grass species by photosynthetic pathway overlooks variation in several functional traits, particularly for C4species. These results indicate that further assessment of lineage‐based differences at other sites and across other grass species distributions may improve representation of C4species in trait comparison analyses and modeling investigations.

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  5. Adams, Henry (Ed.)

    The ubiquity of woody plant expansion across many rangelands globally has led to the hypothesis that the global rise in atmospheric carbon dioxide concentration ([CO2]) is a global driver facilitating C3 woody plant expansion. Increasing [CO2] also influences precipitation patterns seasonally and across the landscape, which often results in the prevalence of drought in rangelands. To test the potential for [CO2] to facilitate woody plant growth, we conducted a greenhouse study for 150 days to measure CO2 effects on juveniles from four woody species (Cornus drummondii C.A. Mey., Rhus glabra L., Gleditsia triacanthos L., Juniperus osteosperma Torr.) that are actively expanding into rangelands of North America. We assessed chronic water-stress (nested within CO2 treatments) and its interaction with elevated [CO2] (800 p.p.m.) on plant growth physiology for 84 days. We measured leaf-level gas exchange, tissue-specific starch concentrations and biomass. We found that elevated [CO2] increased photosynthetic rates, intrinsic water-use efficiencies and leaf starch concentrations in all woody species but at different rates and concentrations. Elevated [CO2] increased leaf starch levels for C. drummondii, G. triacanthos, J. osteosperma and R. glabra by 90, 39, 68 and 41%, respectively. We also observed that elevated [CO2] ameliorated the physiological effects of chronic water stress for all our juvenile woody species within this study. Elevated [CO2] diminished the impact of water stress on the juvenile plants, potentially alleviating an abiotic limitation to woody plant establishment in rangelands, thus facilitating the expansion of woody plants in the future.

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  6. Cooke, Steven (Ed.)

    In the mid-continental grasslands of North America, climate change is increasing the intensity and frequency of extreme weather events. Increasingly severe storms and prolonged periods of elevated temperatures can impose challenges that adversely affect an individual's condition and, ultimately, survival. However, despite mounting evidence that extreme weather events, such as heavy rain storms, can impose short-term physiological challenges, we know little regarding the putative costs of such weather events. To determine the consequences of extreme weather for small endotherms, we tested predictions of the relationships between both severe precipitation events and wet bulb temperatures (an index that combines temperature and humidity) prior to capture with body composition and hematocrit of grasshopper sparrows (Ammodramus savannarum) caught during the breeding season at the Konza Prairie Biological Station, Kansas, USA, between 2014 and 2016. We measured each individual's fat mass, lean mass and total body water using quantitative magnetic resonance in addition to their hematocrit. Individuals exposed to storms in the 24 hours prior to capture had less fat reserves, more lean mass, more water and higher hematocrit than those exposed to moderate weather conditions. Furthermore, individuals stored more fat if they experienced high wet bulb temperatures in the week prior to capture. Overall, the analysis of these data indicate that extreme weather events take a physiological toll on small endotherms, and individuals may be forced to deplete fat stores and increase erythropoiesis to meet the physiological demands associated with surviving a storm. Elucidating the potential strategies used to cope with severe weather may enable us to understand the energetic consequences of increasingly severe weather in a changing world.

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  7. Abstract

    A frequently expressed viewpoint across the Earth science community is that global soil moisture estimates from satellite L‐band (1.4 GHz) measurements represent moisture only in a shallow surface layer (0–5 cm) and consequently are of limited value for studying global terrestrial ecosystems because plants use water from deeper rootzones. Using this argumentation, many observation‐based land surface studies avoid satellite‐observed soil moisture. Here, based on peer‐reviewed literature across several fields, we argue that such a viewpoint is overly limiting for two reasons. First, microwave soil emission depth considerations and statistical considerations of vertically correlated soil moisture information together indicate that L‐band measurements carry information about soil moisture extending below the commonly referenced 5 cm in many conditions. However, spatial variations of effective depths of representation remain uncertain. Second, in reviewing isotopic tracer field studies of plant water uptake, we find a prevalence of vegetation that primarily draws moisture from these upper soil layers. This is especially true for grasslands and croplands covering more than a third of global vegetated surfaces. Even some deeper‐rooted species (i.e., shrubs and trees) preferentially or seasonally draw water from the upper soil layers. Therefore, L‐band satellite soil moisture estimates are more relevant to global vegetation water uptake than commonly appreciated (i.e., relevant beyond only shallow soil processes like soil evaporation). Our commentary encourages the application of satellite soil moisture across a broader range of terrestrial hydrosphere and biosphere studies while urging more rigorous estimates of its effective depth of representation.

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  8. Abstract

    Despite asymmetric competition and a wide array of functional similarities, two ecologically important C4perennial grasses,Andropogon gerardiiandSorghastrum nutans, frequently codominate areas of the mesic tallgrass prairie of the US Great Plains. A subtle difference in their vegetative reproduction strategies may play a role in preventing the exclusion ofS. nutans, the presumed weaker competitor in such regions.

    WhileA. gerardiivegetative tiller densities peak in the early growing season and decline thereafter (determinate recruitment), those ofS. nutansmay continue to increase throughout the growing season (indeterminate recruitment), providing a potential avenue for recovery from more intensive early season competition. However, until now these patterns have only been informally observed in the field.

    We examined the year‐to‐year consistency of growing season vegetative tiller dynamics (measured as seasonal change in tiller densities) of each grass species from an intact tallgrass prairie in Kansas – a site within the core of both species' distributions – over a period of 8 years. Then, to investigate environmental effects on these dynamics, we examined whether they differ across a Kansas landscape varying in topography, fire management regimes, and the abundances of the study species. Finally, we expanded the investigation of environmental effects on growing season tiller dynamics by observing them at the periphery of the species' distributions in central Colorado, where climatic conditions are dryer and the study species' abundances are reduced.

    Synthesis. We found that the tiller densities ofA. gerardiidecline within seasons with striking consistency regardless of spatio‐temporal scale or environmental factors (topography and fire regimes). In contrast, we found the seasonal dynamics ofS. nutanstiller densities were dependent on environmental factors, with seasonal tiller density increases occurring only within the Kansas populations but not consistent between years. These observations lay the groundwork for establishing differences in tiller recruitment determinacy as a potentially important yet underappreciated mechanism for promoting coexistence and codominance among perennial plant species.

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    Free, publicly-accessible full text available March 1, 2024
  9. Abstract

    Plant community assembly outcomes can be contingent upon establishment year (year effects) due to variations in the environment. Stochastic events such as interannual variability in climate, particularly in the first year of community assembly, contribute to unpredictable community outcomes over the short term, but less is known about whether year effects produce transient or persistent states on a decadal timescale. To test for short‐term (5‐year) and persistent (decadal) effects of establishment year climate on community assembly outcomes, we restored prairie in an agricultural field using the same methods in four different years (2010, 2012, 2014, and 2016) that captured a wide range of initial (planting) year climate conditions. Species composition was measured for 5 years in all four restored prairies and for 9 and 11 years in the two oldest restored prairies established under average precipitation and extreme drought conditions. The composition of the four assembled communities showed large and significant differences in the first year of restoration, followed by dynamic change over time along a similar trajectory due to a temporary flush of annual volunteer species. Sown perennial species eventually came to dominate all communities, but communities remained distinct from each other in year five. Precipitation in June and July of the establishment year explained short‐term coarse community metrics (i.e., species richness and grass/forb cover), with wet establishment years resulting in a higher cover of grasses and dry establishment years resulting in a higher cover of forbs in restored communities. Short‐term differences in community composition, species richness, and grass/forb cover in restorations established under average precipitation and drought conditions persisted for 9–11 years, with low interannual variability in the composition of each prairie over the long term, indicating persistently different states on a decadal timescale. Thus, year effects resulting from stochastic variation in climate can have decadal effects on community assembly outcomes.

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  10. Abstract

    Many exogenous factors may influence demographic rates (i.e., births, deaths, immigration, emigration), particularly for migratory birds that must cope with variable weather and habitat throughout their range and annual cycle. In midcontinental grasslands, disturbance (e.g., fire and grazing) and precipitation influence variation in grassland structure and function, but we know little about when and why precipitation is associated with grassland species' vital rates. We related estimates of detection, survival, and emigration toa priorisets of precipitation metrics to test the putative alternative factors influencing movement and mortality in grasshopper sparrows (Ammodramus savannarum). This species is a migratory songbird that exhibits exceptionally high rates of within‐season and between‐season dispersal. Between 2013 and 2020, we captured and resighted grasshopper sparrows in northeastern Kansas, USA, compiling capture histories for 1,332 adult males. We tested predictions of climatic hypotheses explaining variation in survival and emigration throughout a grasshopper sparrow's annual cycle; both survival and emigration were associated with the El Niño‐Southern Oscillation precipitation index (ESPI). Survival was positively related with ESPI during winter, and temporary emigration was curvilinearly related to breeding season ESPI lagged 2 years, with the highest site fidelity associated with intermediate rainfall values. The relationship between rainfall and temporary emigration likely reflects the influence of weather over multiple years on vegetation structure with consequent effects on local demography. This study provides compelling support for the idea that grassland species respond to high interannual variability by adopting dispersal strategies unlike those of many well‐studied migrant birds. Furthermore, the results imply that the consequences of increasing climatic extremes may not be immediately apparent, with demographic consequences lasting for at least a few years.

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