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Research Article

Conditional vulnerability of plant diversity to atmospheric nitrogen deposition across the United States

Samuel M. Simkin, Edith B. Allen, William D. Bowman, Christopher M. Clark, Jayne Belnap, Matthew L. Brooks, Brian S. Cade, View ORCID ProfileScott L. Collins, Linda H. Geiser, Frank S. Gilliam, Sarah E. Jovan, Linda H. Pardo, Bethany K. Schulz, Carly J. Stevens, Katharine N. Suding, Heather L. Throop, and Donald M. Waller
  1. aInstitute of Arctic and Alpine Research and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309;
  2. bDepartment of Botany and Plant Sciences, University of California, Riverside, CA 92521;
  3. cCenter for Conservation Biology, University of California, Riverside, CA 92521;
  4. dNational Center for Environmental Assessment, United States Environmental Protection Agency, Washington, DC 20460;
  5. eSouthwest Biological Science Center, United States Geological Survey, Moab, UT 84532;
  6. fWestern Ecological Research Center, United States Geological Survey, Oakhurst, CA 93644;
  7. gFort Collins Science Center, United States Geological Survey, Fort Collins, CO 80226;
  8. hDepartment of Biology, University of New Mexico, Albuquerque, NM 87131;
  9. iPacific Northwest Region Air Resource Management Program, United States Department of Agriculture Forest Service, Corvallis, OR 97339;
  10. jDepartment of Biological Sciences, Marshall University, Huntington, WV 25755;
  11. kForest Inventory and Analysis Program, United States Department of Agriculture Forest Service, Portland, OR 97339;
  12. lNorthern Research Station, United States Department of Agriculture Forest Service, Burlington, VT 05405;
  13. mForest Inventory and Analysis Program, United States Department of Agriculture Forest Service, Anchorage, AK 99501;
  14. nLancaster Environment Centre, Lancaster University, Lancaster LA1 4YQ, United Kingdom;
  15. oSchool of Earth and Space Exploration, Arizona State University, Tempe, AZ 85287;
  16. pSchool of Life Sciences, Arizona State University, Tempe, AZ 85287;
  17. qDepartment of Botany, University of Wisconsin, Madison, WI 53706

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PNAS April 12, 2016 113 (15) 4086-4091; first published March 28, 2016; https://doi.org/10.1073/pnas.1515241113
Samuel M. Simkin
aInstitute of Arctic and Alpine Research and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309;
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  • For correspondence: samuel.simkin@colorado.edu
Edith B. Allen
bDepartment of Botany and Plant Sciences, University of California, Riverside, CA 92521;
cCenter for Conservation Biology, University of California, Riverside, CA 92521;
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William D. Bowman
aInstitute of Arctic and Alpine Research and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309;
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Christopher M. Clark
dNational Center for Environmental Assessment, United States Environmental Protection Agency, Washington, DC 20460;
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Jayne Belnap
eSouthwest Biological Science Center, United States Geological Survey, Moab, UT 84532;
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Matthew L. Brooks
fWestern Ecological Research Center, United States Geological Survey, Oakhurst, CA 93644;
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Brian S. Cade
gFort Collins Science Center, United States Geological Survey, Fort Collins, CO 80226;
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Scott L. Collins
hDepartment of Biology, University of New Mexico, Albuquerque, NM 87131;
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Linda H. Geiser
iPacific Northwest Region Air Resource Management Program, United States Department of Agriculture Forest Service, Corvallis, OR 97339;
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Frank S. Gilliam
jDepartment of Biological Sciences, Marshall University, Huntington, WV 25755;
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Sarah E. Jovan
kForest Inventory and Analysis Program, United States Department of Agriculture Forest Service, Portland, OR 97339;
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Linda H. Pardo
lNorthern Research Station, United States Department of Agriculture Forest Service, Burlington, VT 05405;
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Bethany K. Schulz
mForest Inventory and Analysis Program, United States Department of Agriculture Forest Service, Anchorage, AK 99501;
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Carly J. Stevens
nLancaster Environment Centre, Lancaster University, Lancaster LA1 4YQ, United Kingdom;
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Katharine N. Suding
aInstitute of Arctic and Alpine Research and Department of Ecology and Evolutionary Biology, University of Colorado, Boulder, CO 80309;
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Heather L. Throop
oSchool of Earth and Space Exploration, Arizona State University, Tempe, AZ 85287;
pSchool of Life Sciences, Arizona State University, Tempe, AZ 85287;
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Donald M. Waller
qDepartment of Botany, University of Wisconsin, Madison, WI 53706
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  1. Edited by Sarah E. Hobbie, University of Minnesota, Saint Paul, Saint Paul, MN, and approved February 23, 2016 (received for review August 4, 2015)

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  • Fig. 1.
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    Fig. 1.

    Herbaceous plant species richness relationships with N deposition. Raw data points (n = 15,136 sites) are gray. Surface plots represent 0.1 (red), 0.5 (median; black), and 0.9 (blue) quantile regression models (median parameters in Table 1) fitted to 3,317 open sites (combined grassland, shrubland, and woodland) (A) and 11,819 closed canopy sites (combined deciduous, evergreen, and mixed forests) (B), as influenced by soil pH. Asterisks indicate significant interactions (P < 0.05).

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    Fig. S1.

    Herbaceous plant species richness relationships with N deposition, as moderated by temperature and precipitation. Raw data points (n = 15,136 sites) are gray. Surface plots represent 0.1 (red), 0.5 (median; black), and 0.9 (blue) quantile regression models (median parameters in Table 1) fitted to 3,317 open sites (combined grassland, shrubland, and woodland) (A and B) and 11,819 closed canopy sites (combined deciduous, evergreen, and mixed forests) (C and D), as influenced by temperature (A and C) and precipitation (B and D). Asterisks indicate significant interactions (P < 0.05). This figure is the counterpart to the pH panels in Fig. 1.

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    Fig. 2.

    Nitrogen deposition (gray-scale) and critical loads for nitrogen deposition based on total graminoid plus forb species richness (colored symbols). The 3,317 open sites (combined grassland, shrubland, and woodland vegetation types) are portrayed with triangles, and the 11,819 closed canopy sites (deciduous, evergreen, and mixed forests) are portrayed with circles. Background deposition values are the average of 27 y of wet deposition (NADP 1985–2011) plus the average of 10 y of dry deposition (CMAQ 2002–2011). Other variation in critical loads is due to the other predictor variables (soil pH, temperature, and precipitation).

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    Fig. S2.

    Critical loads for N deposition at open vegetation sites only, based on total graminoid plus forb species richness. This figure is the same as Fig. 2, except that only the 3,317 open vegetation sites (combined grassland, shrubland, and woodland vegetation types) are portrayed here, so that those sites and the underlying N deposition can be seen more clearly. Background deposition values are the average of 27 y of wet deposition (NADP 1985–2011) plus the average of 10 y of dry deposition (CMAQ 2002–2011).

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    Fig. S3.

    Uncertainty in the critical loads for nitrogen deposition. The percent error in the critical load for each site in Fig. 2 is calculated at each site as the average of the absolute values of the lower and upper endpoints (equivalent to a plus/minus percent error for a symmetric CI) of the 95% confidence interval of 10,000 Monte Carlo simulations of coefficient uncertainty at each site. Background deposition values are the average of 27 y of wet deposition (NADP 1985–2011) plus the average of 10 y of dry deposition (CMAQ 2002–2011).

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    Fig. S4.

    Critical loads (and their 95% confidence interval) of N deposition as a function of soil pH. The 11,819 closed canopy sites (deciduous, evergreen, and mixed forests) are in A, and the 3,317 open sites (combined grassland, shrubland, and woodland vegetation types) are in B.

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    Fig. S5.

    Exceedances of critical loads for nitrogen deposition based on total graminoid plus forb species richness. Exceedances were calculated by subtracting critical loads (of median point estimates) from N deposition values at each site, so negative values indicate that N deposition has been below the critical load and positive values indicate that N deposition has exceeded the critical load. The 3,317 open sites (combined grassland, shrubland, and woodland vegetation types) are portrayed with triangles, and the 11,819 closed canopy sites (deciduous, evergreen, and mixed forests) are portrayed with circles. Background deposition values are the average of 27 y of wet deposition (NADP 1985–2011) plus the average of 10 y of dry deposition (CMAQ 2002–2011).

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    Fig. 3.

    Summary of relationships between plant species richness and N deposition in 46 gradients. Gradients (uniquely defined by vegetation type and data source) contain 6,807 sites, conditional on soil pH, average annual temperature, annual precipitation, and N deposition interactions with each of the other three predictors. In conditionally negative or positive gradients, the relationship was either negative or positive, respectively, for more than half of the range of the moderating variable(s).

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    Fig. 4.

    Magnitude of plant species richness changes associated with N deposition, as moderated individually by (A) soil pH, (B) average temperature, or (C) annual precipitation. Each point, symbolized by the mean N deposition of that gradient (kg⋅ha−1⋅y−1), represents an individual gradient with a single narrow vegetation type. Species richness change is calculated as the simple slope of nitrogen deposition from multiple regression coefficients: βN + (βN*M × Mi), where βN is the parameter for N deposition, βN*M is the parameter for the interaction of N deposition and the moderating variable M, and Mi are the mean (symbol) and range (lines) of the moderating variable M across the gradient. Unlike in Fig. 3, each predictor variable is considered separately.

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    Table S1.

    Summary of vegetation data sources for national surface analysis

    Data nameLocationSourceSites
    AlvarNortheastern USPeet RK, Lee MT, Jennings MD, Faber-Langendoen D. Long database report: VegBank—A permanent, open-access archive for vegetation-plot data. Pages 233–241 in ref. 37. Downloaded from VegBank.36
    COWestern plains of US (Pawnee Nat. Grassland)Peet RK, Lee MT, Jennings MD, Faber-Langendoen D. Long Database Report: VegBank—A permanent, open-access archive for vegetation-plot data. Pages 233–241 in ref. 37. Downloaded from VegBank.17
    CVSSoutheastern USPeet RK, Lee MT, Boyle MF, Wentworth, TR, Schafale, MP, Weakley AS. Long database report: Vegetation-plot database of the Carolina Vegetation Survey. Pages 243–253 in ref. 37. Provided by R. K. Peet and M. T. Lee in 2013 and now available in the Ecological Society of America VegBank.2,611
    FIAContiguous USSchulz BK, Dobelbower K. Short database report: FIADB vegetation diversity and structure indicator (VEG). Page 436 in ref. 37. Provided by B. K. Schulz. Available from apps.fs.fed.us/fiadb-downloads/datamart.html.1,280
    KnutsonIntermountain West of USsagemap.wr.usgs.gov/ESR_Chrono.aspx and ref. 38. Provided by K. C. Knutson.160
    MNUpper Midwest USProvided by Minnesota Biological Survey. Copyright 2013 State of Minnesota, Department of Natural Resources.2,896
    NYNortheastern US (NY Natural Heritage Program)Peet RK, Lee MT, Jennings MD, Faber-Langendoen D. Long database report: VegBank—A permanent, open-access archive for vegetation-plot data. Pages 233–241 in ref. 37. Downloaded from VegBank.75
    PNWPacific Northwest of USPeet RK, Lee MT, Jennings MD, Faber-Langendoen D. Long database report: VegBank—A permanent, open-access archive for vegetation-plot data. Pages 233–241 in ref. 37. Downloaded from VegBank.3,570
    SWSouthwestern USProvided by the Southern Colorado Plateau Network of the US National Park Service and the laboratory of W. D. Bowman.106
    VASoutheastern USProvided by the Virginia Department of Conservation and Recreation, Division of Natural Heritage, VA Plots, the DCR-DNH Vegetation Plots Database. Data exported on March 8, 2013. Now available in VegBank.2,777
    WIUpper Midwest USWaller DM, Amatangelo KL, Johnson S, Rogers DA. Long database report: Wisconsin Vegetation Database—Plant community survey and resurvey data from the Wisconsin Plant Ecology Laboratory. Pages 255–264 in ref. 37. Provided by D. M. Waller.112
    WVSoutheastern USVanderhorst JP, Byers EA, Streets BP. Short database report: Natural Heritage Vegetation Database for West Virginia. Page 440 in ref. 37. Provided by the West Virginia Natural Heritage Program. Now available in VegBank.1,496
     Overall15,136
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    Table S2.

    Summary of predictor variable sources

    VariableSource
    Wet N depositionNADP (2013) Annual National Trends Network (NTN) Maps by Analyte. Available at nadp.sws.uiuc.edu/ntn/maps.aspx (National Atmospheric Deposition Program, Illinois State Water Survey).
    Dry N depositionCMAQ (Community Multiscale Air Quality) model. Provided by R. L. Dennis in 2014 (39).
    Precipitation, mmPRISM (2013) 30-year normals. Parameter-elevation relationships on independent slopes model. Available at prism.nacse.org/normals/ (Northwest Alliance for Computational Science & Engineering at Oregon State University) (1981–2010 normals) (40).
    Temperature, °CPRISM (2013) 30-year normals. Parameter-elevation relationships on independent slopes model. Available at prism.nacse.org/normals/ (Northwest Alliance for Computational Science & Engineering at Oregon State University) (1981–2010 normals) (40).
    Soil pHProvided with vegetation data; otherwise from USDA SSURGO soil dataset. SSURGO (2014) Geospatial Data Gateway. Available at https://gdg.sc.egov.usda.gov/GDGOrder.aspx?order=QuickState (USDA Natural Resources Conservation Service).
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    Table S3.

    Vegetation Alliances with sufficient data for gradient analyses

    Vegetation Alliance (33) (from National Vegetation Classification)N deposition, kg⋅ha−1⋅y−1Sites
    Min.Max.
    Abies lasiocarpa-Picea engelmannii forest (ABLA)1.75.0355
    Acer saccharum-Betula alleghaniensis-(Fagus grandifolia) forest (ACSA)4.115.2399
    Acer saccharum-Tilia americana-(Quercus rubra) forest (ACETIL)4.717.0479
    Andropogon gerardii-(Sorghastrum nutans) herbaceous (ANGE)8.714.064
    Artemisia tridentata shrubland (ARTR)1.34.6314
    Betula papyrifera forest (BEPA)3.411.425
    Fagus grandifolia-Quercus rubra-Quercus alba forest (FAGR)7.014.926
    Festuca idahoensis alpine herbaceous (FEID)1.84.6330
    Liriodendron tulipifera forest (LITU)8.415.620
    Pinus contorta forest (PICO)1.74.8603
    Pinus palustris woodland (PIPA)5.615.1199
    Pinus ponderosa forest (PIPO)1.56.0238
    Pinus strobus forest (PIST)4.113.950
    Pinus taeda forest (PITA)6.614.4206
    Populus tremuloides forest (POTR)1.312.3362
    Pseudotsuga menziesii forest (PSME)1.47.5333
    Pseudotsuga menziesii giant forest (PSME)1.43.6106
    Pseudotsuga menziesii woodland (PSME)1.54.4656
    Quercus alba-(Quercus rubra, Carya spp.) forest (QUAL)6.617.81,362
    Quercus prinus-Quercus (alba, falcata, rubra, velutina) forest (QUPR)9.315.223
    Quercus spp.-Pinus (rigida, echinata) forest (QUEPIN)7.917.9233
    Quercus virginiana-(Sabal palmetto) forest (QUEVIR)6.012.821
    Schizachyrium scoparium-Bouteloua curtipendula herbaceous (SCSC)8.414.3267
    Symphoricarpos albus shrubland (SYAL)1.64.131
    Thuja occidentalis forest (THOC)4.29.124
    Tilia americana-Fraxinus americana-(Acer saccharum) woodland (TIAM)4.714.181
     Overall1.317.96,807
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    Table 1.

    Parameter coefficients for species richness from median quantile regressions

    NameOpen canopy (±1 SE)Closed canopy (±1 SE)
    Intercept14.9 (3.42)*13.6 (2.55)*
    N4.69 (0.60)*0.449 (0.33)n
    N2−0.494 (0.02)*−0.125 (0.01)*
    pH−2.17 (0.46)*−1.49 (0.37)*
    Precip−0.011 (0.002)*−0.003 (0.001)*
    Temp−0.059 (0.18)n−0.321 (0.04)*
    N:pH0.475 (0.07)*0.543 (0.04)*
    N:precip0.002 (0.001)*NA
    N:temp−0.073 (0.03)+NA
    • Regressions represent herbaceous plant species richness response to N deposition (kg⋅ha−1⋅y−1; quadratic), soil pH, total annual precipitation (mm), average annual temperature (°C), and interactions of N (deposition) with pH, precipitation, and temperature. An NA (not applicable) indicates that term didn't appear in best model. Sample size is 11,819 sites for closed canopy (deciduous forest, evergreen forest, and mixed forest) and 3,317 sites for open canopy (grassland, shrubland, and woodland). Level of significance is indicated as follows: nP ≥ 0.05, +P < 0.05, or *P < 0.001.

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    Table 2.

    Critical loads (CLs) of N deposition for herbaceous plant species richness

    VegetationCL expression (partial derivative of species richness equation)CL (kg N⋅ha−1⋅y−1)CL error§
    Mean†Range†Range of 95% CI‡
    Open canopy vegetation[4.690 + (0.475 · (soil pH)) + (0.0018 · (mm of precip.)) + (−0.073 · (temp. (°C)))]/(−2 · −0.494)8.77.4–10.36.4–11.3−4.5%, 4.8%
    Closed canopy vegetation[0.449 + (0.543 · (soil pH))]/(−2 · −0.125)13.47.9–19.66.8–22.2−6.2%, 7.7%
    • The critical load (CL) expression is derived using the partial derivative with respect to nitrogen of the species richness equation in Table 1, and then evaluated locally with site-specific soil pH, precipitation, and temperature values.

    • ↵† Mean and range of CLs across sites, reflecting variation in soil pH, precipitation, and temperature variables across sites but not uncertainty in coefficient estimates.

    • ↵‡ Range of CL 95% confidence interval endpoints across sites (Fig. S3), reflecting both ecological variability (soil pH and climate variables) and uncertainty in coefficient estimates, with the latter calculated from the 2.5th and 97.5th percentiles of 10,000 Monte Carlo simulations of coefficient uncertainty.

    • ↵§ Average of the site-specific CL % errors, calculated from the lower and upper endpoints of the 95% confidence interval of Monte Carlo simulations of coefficient uncertainty repeated at each site.

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    Table S4.

    Critical loads (CLs) of N deposition for level 1 ecoregions in the present study compared with previously cited estimates in Pardo et al. (18)

    Level 1 ecoregionCL of N dep. cited in table 5 of ref. 18 (Pardo et al.), kg⋅ha−1⋅y−1CL of N dep. (total herbaceous richness) from present study, kg⋅ha−1⋅y−1Compared with Pardo et al. (18)
    Northern forests7–21 (hardwood forest alteration of herbaceous understory)Open: 8.0–9.8 (mean = 8.9, n = 75)New
    Closed: 8.0–18.9 (mean = 13.8, n = 1,955)Similar
    Northwestern forested mountains4–10 (alpine grassland species composition change)Open: 8.0–10.2 (mean = 9.1, n = 1,429)Similar
    Closed: 10.8–19.6 (mean = 15.3, n = 2,113)New
    Marine west coast forestsNo vascular plant CLOpen: No data
    Closed: 10.4–15.0 (mean = 12.8, n = 24)New
    Eastern temperate forests<17.5 (hardwood forest declines in species-rich genera)Open: 6.6–9.7 (mean = 7.9, n = 947)New
    Closed: 7.8–19.3 (mean = 12.5, n = 7,378)Lower
    Great plains5–15 (tallgrass prairie community shifts)Open: 8.3–9.8 (mean = 9.3, n = 618)Similar
    Closed: 11.3–19.6 (mean = 16.6, n = 274)New
    North American desert3–8.4 (warm desert decrease of native forbs)Open: 8.3–9.9 (mean = 9.2, n = 240)Higher
    Closed: 13.5–17.0 (mean = 16.5, n = 32)New
    Temperate SierrasNo vascular plant CLOpen: 8.6–8.7 (mean = 8.65, n = 3)New
    Closed: 14.8–14.8 (mean = 14.8, n = 42)New
    • Critical load estimates in the present study are calculated using the two equations in Table 2. “Open” refers to open canopy vegetation whereas “Closed” refers to closed canopy forest vegetation. We did not have any vegetation sites with critical loads to summarize for the remaining two ecoregions: namely the wet tropical forest and the Mediterranean California ecoregions.

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    Table S5.

    Summary of herbaceous plant species richness responses to N deposition in gradients

    Gradient codeN deposition responseData sourceNVC AllianceMajor vegetation typeModelR2WeightSitesN_dep minN_dep meanN_dep maxPrecip mm minPrecip mm meanPrecip mm maxTemp °C minTemp °C meanTemp °C maxpH minpH meanpH max
    ACSA2Spp decline unconditionally with N depCVSAcer saccharum-Betula alleghaniensis-(Fagus grandifolia) forestBroad-leaved forestN_dep + pH + temp_C + 10.2690.73349.1211.9813.771,2031,6402,3839.011.015.24.234.875.80
    ACSA3No N dep responseFIAAcer saccharum-Betula alleghaniensis-(Fagus grandifolia) forestBroad-leaved forestpH + 10.1070.621764.128.3014.536801,0341,6132.95.68.93.424.496.54
    ACSA4No N dep responseMNAcer saccharum-Betula alleghaniensis-(Fagus grandifolia) forestBroad-leaved forest10.0000.42234.686.928.756256817903.84.35.15.006.026.70
    ACSA5Spp decline conditional on precipWVAcer saccharum-Betula alleghaniensis-(Fagus grandifolia) forestBroad-leaved forestN_dep + N:precip + pH + precip_mm + temp_C + 10.4260.651669.0811.7915.238861,2171,7186.38.812.93.204.487.90
    ACETIL1No N dep responseFIAAcer saccharum-Tilia americana-(Quercus rubra) forestBroad-leaved forest10.0000.38324.939.2712.557109631,3394.16.18.63.834.966.46
    ACETIL2Spp decline conditional on precip and tempMNAcer saccharum-Tilia americana-(Quercus rubra) forestBroad-leaved forestN_dep + N:precip + N:temp + pH + precip_mm + temp_C + 10.3170.313524.698.9214.406077418953.15.58.24.906.167.90
    ACETIL3Spp increase conditional on tempWIAcer saccharum-Tilia americana-(Quercus rubra) forestBroad-leaved forestN_dep + N:temp + temp_C + 10.3001.00956.0210.7517.047288489454.27.18.94.305.907.40
    BEPANo N dep responseFIABetula papyrifera forestBroad-leaved forestpH + precip_mm + 10.2110.41253.376.3311.426649631,5013.04.87.93.414.926.49
    FAGRNo N dep responseVAFagus grandifolia-Quercus rubra-Quercus alba forestBroad-leaved forestpH + 10.2650.42266.9811.6114.941,0541,1421,24313.214.415.83.804.455.40
    LITUSpp decline unconditionally with N depFIALiriodendron tulipifera forestBroad-leaved forestN_dep + precip_mm + 10.4891.00208.4012.7215.629851,1341,3308.711.414.43.945.056.00
    POTR1No N dep responseFIAPopulus tremuloides forestBroad-leaved forestpH + temp_C + 10.2180.25854.897.4512.316277791,1592.84.97.63.885.047.04
    POTR2Spp decline conditional on precipMNPopulus tremuloides forestBroad-leaved forestN_dep + N:precip + precip_mm + 10.0770.312274.696.979.295496557823.04.15.34.806.287.90
    POTR3No N dep responsePNWPopulus tremuloides forestBroad-leaved forestprecip_mm + temp_C + 10.4460.67501.302.514.073256231,3591.14.47.85.506.547.40
    QUAL1Spp decline conditional on pHCVSQuercus alba-(Quercus rubra, Carya spp.) forestBroad-leaved forestN_dep + N:pH + pH + precip_mm + temp_C + 10.4200.341829.5411.1215.571,1151,3652,1719.213.117.33.504.916.54
    QUAL2No N dep responseFIAQuercus alba-(Quercus rubra, Carya spp.) forestBroad-leaved forestpH + 10.0120.252026.6311.2617.847151,0661,4483.810.714.63.495.157.82
    QUAL3No N dep responseMNQuercus alba-(Quercus rubra, Carya spp.) forestBroad-leaved forestpH + 10.0850.33576.8311.0413.316258178924.06.98.35.406.457.50
    QUAL4No N dep responseVAQuercus alba-(Quercus rubra, Carya spp.) forestBroad-leaved forestpH + temp_C + 10.2900.443557.0810.7615.389441,1751,7117.112.015.73.004.717.90
    QUAL5Spp decline unconditionally with N depWIQuercus alba-(Quercus rubra, Carya spp.) forestBroad-leaved forestN_dep + precip_mm + temp_C + 10.2980.194510.1912.2716.207898859267.08.08.84.706.027.10
    QUAL6No N dep responseWVQuercus alba-(Quercus rubra, Carya spp.) forestBroad-leaved forestpH + precip_mm + 10.2730.455218.4511.3015.238741,1631,7256.510.513.03.304.477.50
    QUPRSpp decline unconditionally with N depFIAQuercus prinus-Quercus (alba, falcata, rubra, velutina) forestBroad-leaved forestN_dep + 10.2140.25239.2711.8215.199111,1291,3387.910.413.93.234.215.36
    QUEPIN2Spp decline conditional on temp precip and pHCVSQuercus spp.-Pinus (rigida, echinata) forestMixed forestN_dep + N:pH + N:precip + N:temp + pH + precip_mm + temp_C + 10.5351.001207.8811.1417.901,0851,2981,5019.714.219.73.184.485.74
    QUEPIN1No N dep responseVAQuercus spp.-Pinus (rigida, echinata) forestMixed forestpH + temp_C + 10.3310.561138.4010.9513.739801,1541,6577.412.615.53.304.608.00
    QUEVIRNo N dep responseCVSQuercus virginiana-(Sabal palmetto) forestMixed forest10.0000.63216.019.3512.751,1851,2651,45117.318.619.24.565.707.85
    ABLASpp increase conditional on precip and tempPNWAbies lasiocarpa-Picea engelmannii forestConifer forestN_dep + N:precip + N:temp + precip_mm + temp_C + 10.3310.423551.682.995.004171,1192,133−1.52.77.44.806.097.60
    PICOSpp increase conditional on temp and pHPNWPinus contorta forestConifer forestN_dep + N:pH + N:temp + pH + precip_mm + temp_C + 10.2260.726031.692.974.773387731,565−1.42.97.44.806.238.20
    PIPO1No N dep responseFIAPinus ponderosa forestConifer foresttemp_C + 10.0950.33281.602.936.003356902,1313.66.59.24.606.117.61
    PIPO2No N dep responsePNWPinus ponderosa forestConifer foresttemp_C + 10.1090.542101.462.364.353286401,5540.86.59.15.006.407.60
    PIST1No N dep responseFIAPinus strobus forestConifer forestpH + temp_C + 10.3600.45254.097.0910.696709241,2573.15.57.53.784.705.82
    PIST2No N dep responseWVPinus strobus forestConifer forestpH + 10.2351.00258.9410.5913.909701,1431,4647.010.212.23.704.606.50
    PITA1Spp decline conditional on temp and pHCVSPinus taeda forestConifer forestN_dep + N:pH + N:temp + pH + temp_C + 10.5840.711826.5610.9914.351,0831,2811,48913.716.822.63.004.447.20
    PITA2No N dep responseVAPinus taeda forestConifer forestprecip_mm + temp_C + 10.2680.26247.3811.5414.411,0641,0921,12414.514.815.34.604.885.30
    PSME1Spp decline unconditionally with N depFIAPseudotsuga menziesii forestConifer forestN_dep + 10.0880.71621.423.557.513971,6094,3464.78.812.64.005.567.43
    PSME2Spp increase conditional on temp and pHPNWPseudotsuga menziesii forestConifer forestN_dep + N:pH + N:temp + pH + precip_mm + temp_C + 10.1590.482711.612.784.573056111,988−1.33.98.74.806.498.20
    PSME3Spp increase unconditionally with N depPNWPseudotsuga menziesii giant forestConifer forestN_dep + N:pH + pH + 10.2500.361061.431.973.623795579193.66.39.65.006.507.00
    THOCNo N dep responseFIAThuja occidentalis forestConifer forestpH + 10.3200.38244.176.819.086178321,2352.74.87.23.695.467.90
    TIAMSpp decline conditional on tempMNTilia americana-Fraxinus americana-(Acer saccharum) woodlandBroad-leaved woodlandN_dep + N:temp + precip_mm + temp_C + 10.6000.40814.6910.3314.065837438863.86.37.75.006.717.90
    PIPA2Spp decline conditional on temp and pHCVSPinus palustris woodlandConifer woodlandN_dep + N:pH + N:temp + pH + precip_mm + temp_C + 10.6140.571995.598.9915.061,1031,3561,69715.218.822.83.404.595.98
    PSME4Spp increase unconditionally with N depPNWPseudotsuga menziesii woodlandConifer woodlandN_dep + N:temp + precip_mm + temp_C + 10.2890.556561.452.434.402446171,5440.34.48.85.006.398.20
    ARTR1Spp decline unconditionally with N depKnutsonArtemisia tridentata shrublandShrublandN_dep + precip_mm + 10.1500.131431.262.093.792052653316.89.010.77.007.989.10
    ARTR2Spp increase conditional on pHPNWArtemisia tridentata shrublandShrublandN_dep + N:pH + pH + precip_mm + temp_C + 10.2290.501711.602.744.572435439850.93.99.65.206.648.20
    SYALNo N dep responsePNWSymphoricarpos albus shrublandShrublandpH + precip_mm + temp_C + 10.5461.00311.602.534.073305138772.55.38.15.206.728.20
    ANGE1Spp decline conditional on tempMNAndropogon gerardii-(Sorghastrum nutans) herbaceousGrasslandN_dep + N:temp + precip_mm + temp_C + 10.3510.67648.7112.0614.016167498826.37.07.85.507.138.20
    FEIDSpp increase conditional on tempPNWFestuca idahoensis Alpine herbaceousGrasslandN_dep + N:temp + pH + precip_mm + temp_C + 10.1730.713301.812.904.573636121,088−0.83.38.85.306.608.20
    SCSCSpp decline unconditionally with N depMNSchizachyrium scoparium-Bouteloua curtipendula herbaceousGrasslandN_dep + temp_C + 10.0930.222678.3711.7014.286047518955.27.08.35.407.227.90

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Conditional vulnerability of plant diversity to atmospheric nitrogen deposition across the United States
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Diversity and nitrogen deposition
Samuel M. Simkin, Edith B. Allen, William D. Bowman, Christopher M. Clark, Jayne Belnap, Matthew L. Brooks, Brian S. Cade, Scott L. Collins, Linda H. Geiser, Frank S. Gilliam, Sarah E. Jovan, Linda H. Pardo, Bethany K. Schulz, Carly J. Stevens, Katharine N. Suding, Heather L. Throop, Donald M. Waller
Proceedings of the National Academy of Sciences Apr 2016, 113 (15) 4086-4091; DOI: 10.1073/pnas.1515241113

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Diversity and nitrogen deposition
Samuel M. Simkin, Edith B. Allen, William D. Bowman, Christopher M. Clark, Jayne Belnap, Matthew L. Brooks, Brian S. Cade, Scott L. Collins, Linda H. Geiser, Frank S. Gilliam, Sarah E. Jovan, Linda H. Pardo, Bethany K. Schulz, Carly J. Stevens, Katharine N. Suding, Heather L. Throop, Donald M. Waller
Proceedings of the National Academy of Sciences Apr 2016, 113 (15) 4086-4091; DOI: 10.1073/pnas.1515241113
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