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Title: A sequence of multiyear wet and dry periods provides opportunities for grass recovery and state change reversals

Multiyear periods (≥4 years) of extreme rainfall are increasing in frequency as climate continues to change, yet there is little understanding of how rainfall amount and heterogeneity in biophysical properties affect state changes in a sequence of wet and dry periods. Our objective was to examine the importance of rainfall periods, their legacies, and vegetation and soil properties to either the persistence of woody plants or a shift toward perennial grass dominance and a state reversal. We examined a 28‐year record of rainfall consisting of a sequence of multiyear periods (average, dry, wet, dry, average) for four ecosystem types in the Jornada Basin. We analyzed relationships between above ground net primary production (ANPP) and rainfall for three plant functional groups that characterize alternative states (perennial grasses, other herbaceous plants, dominant shrubs). A multimodel comparison was used to determine the relative importance of rainfall, soil, and vegetation properties. For perennial grasses, the greatest mean ANPP in mesquite‐ and tarbush‐dominated shrublands occurred in the wet period and in the dry period following the wet period in grasslands. Legacy effects in grasslands were asymmetric, where the lowest production was found in a dry period following an average period, and the greatest production occurred in a dry period following a wet period. For other herbaceous plants, in contrast, the greatest ANPP occurred in the wet period. Mesquite was the only dominant shrub species with a significant positive response in the wet period. Rainfall amount was a poor predictor of ANPP for each functional group when data from all periods were combined. Initial herbaceous biomass at the plant scale, patch‐scale biomass, and soil texture at the landscape scale improved the predictive relationships of ANPP compared with rainfall alone. Under future climate, perennial grass production is expected to benefit the most from wet periods compared with other functional groups with continued high grass production in subsequent dry periods that can shift (desertified) shrublands toward grasslands. The continued dominance by shrubs will depend on the effects that rainfall has on perennial grasses and the sequence of high‐ and low‐rainfall periods rather than the direct effects of rainfall on shrub production.

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Award ID(s):
2025166 1832194
Author(s) / Creator(s):
Publisher / Repository:
Wiley Blackwell (John Wiley & Sons)
Date Published:
Journal Name:
Ecological Monographs
Medium: X
Sponsoring Org:
National Science Foundation
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  1. null (Ed.)
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Volume-weighted mean TDP concentrations in leachate for each crop-year and for the entire 7-year study period were calculated as the total dissolved P leaching flux (kg ha−1) divided by the total drainage (m3 ha−1). One-way ANOVA with time (crop-year) as the fixed factor was conducted to compare total annual drainage rates, P leaching rates, volume-weighted mean TDP concentrations, and maximum aboveground biomass among the cropping systems over all seven crop-years as well as with TDP concentrations from local lakes, streams, and groundwater wells. When a significant (α = 0.05) difference was detected among the groups, we used the Tukey honest significant difference (HSD) post-hoc test to make pairwise comparisons among the groups. In the case of maximum aboveground biomass, we used the Tukey–Kramer method to make pairwise comparisons among the groups because the absence of poplar data after the 2013 harvest resulted in unequal sample sizes. We also used the Tukey–Kramer method to compare the frequency distributions of TDP concentrations in all of the soil leachate samples with concentrations in lakes, streams, and groundwater wells, since each sample category had very different numbers of measurements. Individual spreadsheets in “data table_leaching_dissolved organic carbon and nitrogen.xls” 1.    annual precip_drainage 2.    biomass_corn, perennial grasses 3.    biomass_poplar 4.    annual N leaching _vol-wtd conc 5.    Summary_N leached 6.    annual DOC leachin_vol-wtd conc 7.    growing season length 8.    correlation_nh4 VS no3 9.    correlations_don VS no3_doc VS don Each spreadsheet is described below along with an explanation of variates. Note that ‘nan’ indicate data are missing or not available. First row indicates header; second row indicates units 1. Spreadsheet: annual precip_drainage Description: Precipitation measured from nearby Kellogg Biological Station (KBS) Long Term Ecological Research (LTER) Weather station, over 2009-2016 study period. Data shown in Figure 1; original data source for precipitation ( Drainage estimated from SALUS crop model. Note that drainage is percolation out of the root zone (0-125 cm). Annual precipitation and drainage values shown here are calculated for growing and non-growing crop periods. Variate    Description year    year of the observation crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” precip_G    precipitation during growing period (milliMeter) precip_NG    precipitation during non-growing period (milliMeter) drainage_G    drainage during growing period (milliMeter) drainage_NG    drainage during non-growing period (milliMeter)      2. Spreadsheet: biomass_corn, perennial grasses Description: Maximum aboveground biomass measurements from corn, switchgrass, miscanthus, native grass and restored prairie plots in Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2009-2015. Data shown in Figure 2.   Variate    Description year    year of the observation date    day of the observation (mm/dd/yyyy) crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” replicate    each crop has four replicated plots, R1, R2, R3 and R4 station    stations (S1, S2 and S3) of samplings within the plot. For more details, refer to link ( species    plant species that are rooted within the quadrat during the time of maximum biomass harvest. See protocol for more information, refer to link ( For maize biomass, grain and whole biomass reported in the paper (weed biomass or surface litter are excluded). Surface litter biomass not included in any crops; weed biomass not included in switchgrass and miscanthus, but included in grass mixture and prairie. fraction    Fraction of biomass biomass_plot    biomass per plot on dry-weight basis (Grams_Per_SquareMeter) biomass_ha    biomass (megaGrams_Per_Hectare) by multiplying column biomass per plot with 0.01 3. Spreadsheet: biomass_poplar Description: Maximum aboveground biomass measurements from poplar plots in Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2009-2015. Data shown in Figure 2. Note that poplar biomass was estimated from crop growth curves until the poplar was harvested in the winter of 2013-14. Variate    Description year    year of the observation method    methods of poplar biomass sampling date    day of the observation (mm/dd/yyyy) replicate    each crop has four replicated plots, R1, R2, R3 and R4 diameter_at_ground    poplar diameter (milliMeter) at the ground diameter_at_15cm    poplar diameter (milliMeter) at 15 cm height biomass_tree    biomass per plot (Grams_Per_Tree) biomass_ha    biomass (megaGrams_Per_Hectare) by multiplying biomass per tree with 0.01 4. Spreadsheet: annual N leaching_vol-wtd conc Description: Annual leaching rate (kiloGrams_N_Per_Hectare) and volume-weighted mean N concentrations (milliGrams_N_Per_Liter) of nitrate (no3) and dissolved organic nitrogen (don) in the leachate samples collected from corn, switchgrass, miscanthus, native grass, restored prairie and poplar plots in Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2009-2016. Data for nitrogen leached and volume-wtd mean N concentration shown in Figure 3a and Figure 3b, respectively. Note that ammonium (nh4) concentration were much lower and often undetectable (<0.07 milliGrams_N_Per_Liter). Also note that in 2009 and 2010 crop-years, data from some replicates are missing.    Variate    Description crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” crop-year    year of the observation replicate    each crop has four replicated plots, R1, R2, R3 and R4 no3 leached    annual leaching rates of nitrate (kiloGrams_N_Per_Hectare) don leached    annual leaching rates of don (kiloGrams_N_Per_Hectare) vol-wtd no3 conc.    Volume-weighted mean no3 concentration (milliGrams_N_Per_Liter) vol-wtd don conc.    Volume-weighted mean don concentration (milliGrams_N_Per_Liter) 5. Spreadsheet: summary_N leached Description: Summary of total amount and forms of N leached (kiloGrams_N_Per_Hectare) and the percent of applied N lost to leaching over the seven years for corn, switchgrass, miscanthus, native grass, restored prairie and poplar plots in Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2009-2016. Data for nitrogen amount leached shown in Figure 4a and percent of applied N lost shown in Figure 4b. Note the fraction of unleached N includes in harvest, accumulation in root biomass, soil organic matter or gaseous N emissions were not measured in the study. Variate    Description crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” no3 leached    annual leaching rates of nitrate (kiloGrams_N_Per_Hectare) don leached    annual leaching rates of don (kiloGrams_N_Per_Hectare) N unleached    N unleached (kiloGrams_N_Per_Hectare) in other sources are not studied % of N applied N lost to leaching    % of N applied N lost to leaching 6. Spreadsheet: annual DOC leachin_vol-wtd conc Description: Annual leaching rate (kiloGrams_Per_Hectare) and volume-weighted mean N concentrations (milliGrams_Per_Liter) of dissolved organic carbon (DOC) in the leachate samples collected from corn, switchgrass, miscanthus, native grass, restored prairie and poplar plots in Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2009-2016. Data for DOC leached and volume-wtd mean DOC concentration shown in Figure 5a and Figure 5b, respectively. Note that in 2009 and 2010 crop-years, water samples were not available for DOC measurements.     Variate    Description crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” crop-year    year of the observation replicate    each crop has four replicated plots, R1, R2, R3 and R4 doc leached    annual leaching rates of nitrate (kiloGrams_Per_Hectare) vol-wtd doc conc.    volume-weighted mean doc concentration (milliGrams_Per_Liter) 7. Spreadsheet: growing season length Description: Growing season length (days) of corn, switchgrass, miscanthus, native grass, restored prairie and poplar plots in the Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2009-2015. Date shown in Figure S2. Note that growing season is from the date of planting or emergence to the date of harvest (or leaf senescence in case of poplar).   Variate    Description crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” year    year of the observation growing season length    growing season length (days) 8. Spreadsheet: correlation_nh4 VS no3 Description: Correlation of ammonium (nh4+) and nitrate (no3-) concentrations (milliGrams_N_Per_Liter) in the leachate samples from corn, switchgrass, miscanthus, native grass, restored prairie and poplar plots in Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2013-2015. Data shown in Figure S3. Note that nh4+ concentration in the leachates was very low compared to no3- and don concentration and often undetectable in three crop-years (2013-2015) when measurements are available. Variate    Description crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” date    date of the observation (mm/dd/yyyy) replicate    each crop has four replicated plots, R1, R2, R3 and R4 nh4 conc    nh4 concentration (milliGrams_N_Per_Liter) no3 conc    no3 concentration (milliGrams_N_Per_Liter)   9. Spreadsheet: correlations_don VS no3_doc VS don Description: Correlations of don and nitrate concentrations (milliGrams_N_Per_Liter); and doc (milliGrams_Per_Liter) and don concentrations (milliGrams_N_Per_Liter) in the leachate samples of corn, switchgrass, miscanthus, native grass, restored prairie and poplar plots in Great Lakes Bioenergy Research Center (GLBRC) Biomass Cropping System Experiment (BCSE) during 2013-2015. Data of correlation of don and nitrate concentrations shown in Figure S4 a and doc and don concentrations shown in Figure S4 b. Variate    Description crop    “corn” “switchgrass” “miscanthus” “nativegrass” “restored prairie” “poplar” year    year of the observation don    don concentration (milliGrams_N_Per_Liter) no3     no3 concentration (milliGrams_N_Per_Liter) doc    doc concentration (milliGrams_Per_Liter) 
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  3. Abstract

    In dryland soils, spatiotemporal variation in surface soils (0–10 cm) plays an important role in the function of the “critical zone” that extends from canopy to groundwater. Understanding connections between soil microbes and biogeochemical cycling in surface soils requires repeated multivariate measurements of nutrients, microbial abundance, and microbial function. We examined these processes in resource islands and interspaces over a two‐month period at a Chihuahuan Desert bajada shrubland site. We collected soil inProsopis glandulosa(honey mesquite),Larrea tridentata(creosote bush), and unvegetated (interspace) areas to measure soil nutrient concentrations, microbial biomass, and potential soil enzyme activity. We monitored the dynamics of these belowground processes as soil conditions dried and then rewetted due to rainfall. Most measured variables, including inorganic nutrients, microbial biomass, and soil enzyme activities, were greater under shrubs during both wet and dry periods, with the highest magnitudes under mesquite followed by creosote bush and then interspace. One exception was nitrate, which was highly variable and did not show resource island patterns. Temporally, rainfall pulses were associated with substantial changes in soil nutrient concentrations, though resource island patterns remained consistent during all phases of the soil moisture pulse. Microbial biomass was more consistent than nutrients, decreasing only when soils were driest. Potential enzyme activities were even more consistent and did not decline in dry periods, potentially helping to stimulate observed pulses in CO2efflux following rain events observed at a co‐located eddy flux tower. These results indicate a critical zone with organic matter cycling patterns consistently elevated in shrub resource islands (which varied by shrub species), high decomposition potential that limits soil organic matter accumulation across the landscape, and nitrate fluxes that are decoupled from the organic matter pathways.

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  4. Abstract Aims

    Grassland-to-shrubland transition is a common form of land degradation in drylands worldwide. It is often attributed to changes in disturbance regimes, particularly overgrazing. A myriad of direct and indirect effects (e.g., accelerated soil erosion) of grazing may favor shrubs over grasses, but their relative importance is unclear. We tested the hypothesis that topsoil “winnowing” by wind erosion would differentially affect grass and shrub seedling establishment to promote shrub recruitment over that of grass.


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    Non-winnowed soils were finer-textured and had higher nutrient contents than winnowed soils, but based on desorption curves, winnowed soils had more plant-available moisture. Contrary to expectations, seed germination and seedling growth on winnowed and non-winnowed soils were comparable within a given species. The N2-fixing deciduous shrubP. glandulosawas first to emerge and complete germination, and had the greatest biomass accumulation of all species.


    Germination and early seedling growth of grasses and shrubs on winnowed soils were not adversely nor differentially affected comparing with that observed on non-winnowed soils under well-watered greenhouse conditions. Early germination and rapid growth may giveP. glandulosaa competitive advantage over grasses and other shrub species at the establishment stage in grazed grasslands. Field establishment experiments are needed to confirm our findings in these controlled environment trials.

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  5. Quantifying the relationship of different grass functional groups to increasing woody plant cover is necessary to better understand the effects of woody plant encroachment on grasslands. This study explored biomass production responses of three perennial grass groups based on photosynthetic pathway and potential canopy height (C4 short-grasses, C3 midgrasses, and C4 midgrasses) to different percent canopy covers of the surrounding deciduous woody legume, honey mesquite (Prosopis glandulosa). Two methods were used to determine mesquite canopy cover, line-intercept and geospatial analysis of aerial images, and both were used to predict production of the three grass groups. Five years of grass production data were included in the mesquite cover/grass production regressions. Two yr had extreme grass production responses, one due to drought and the other to high rainfall. Of the 3 remaining yr, best-fit curves were negative linear for C4 short-grasses and C3 midgrasses and negative sigmoidal for C4 midgrasses using both cover determination methods, although slopes of the curves differed between cover determination methods. C4 midgrasses were more sensitive than the other grass groups to increasing mesquite cover. Loss of production potential when mesquite cover increased from 0% to 35% was 75.5%, 28.7%, and 23.2% for C4 midgrasses, C3 midgrasses, and C4 short-grasses, respectively. Moreover, production potential of C4 midgrasses under no mesquite cover was 3 and 6 times greater than C3 midgrasses or C4 short-grasses, respectively. Spatial settings of the different grass groups in relation to mesquite tree size and size of intercanopy areas provided indirect evidence that the process of mesquite encroachment in the past 50−100 yr may have negatively impacted C4 midgrasses more than the other grass groups. Results suggest that gains in grass production following mesquite treatment would be limited if the system has degraded to where only C3 midgrasses and C4 short-grasses dominate. 
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