Despite their low contribution to forest carbon stocks, lianas (woody vines) play an important role in the carbon dynamics of tropical forests. As structural parasites, they hinder tree survival, growth and fecundity; hence, they negatively impact net ecosystem productivity and long‐term carbon sequestration. Competition (for water and light) drives various forest processes and depends on the local abundance of resources over time. However, evaluating the relative role of resource availability on the interactions between lianas and trees from empirical observations is particularly challenging. Previous approaches have used labour‐intensive and ecosystem‐scale manipulation experiments, which are infeasible in most situations. We propose to circumvent this challenge by evaluating the uncertainty of water and light capture processes of a process‐based vegetation model (ED2) including the liana growth form. We further developed the liana plant functional type in ED2 to mechanistically simulate water uptake and transport from roots to leaves, and start the model from prescribed initial conditions. We then used the PEcAn bioinformatics platform to constrain liana parameters and run uncertainty analyses. Baseline runs successfully reproduced ecosystem gas exchange fluxes (gross primary productivity and latent heat) and forest structural features (leaf area index, aboveground biomass) in two sites (Barro Colorado Island, Panama and Paracou, French Guiana) characterized by different rainfall regimes and levels of liana abundance. Model uncertainty analyses revealed that water limitation was the factor driving the competition between trees and lianas at the drier site (BCI), and during the relatively short dry season of the wetter site (Paracou). In young patches, light competition dominated in Paracou but alternated with water competition between the wet and the dry season on BCI according to the model simulations. The modelling workflow also identified key liana traits (photosynthetic quantum efficiency, stomatal regulation parameters, allometric relationships) and processes (water use, respiration, climbing) driving the model uncertainty. They should be considered as priorities for future data acquisition and model development to improve predictions of the carbon dynamics of liana‐infested forests.
Lianas are prevalent in Neotropical forests, where liana‐tree competition can be intense, resulting in reduced tree growth and survival. The ability of lianas to grow relative to trees during the dry season suggests that liana‐tree competition is also strongest in the dry season. If correct, the predicted intensification of the drying trend over large areas of the tropics in the future may therefore intensify liana‐tree competition resulting in a reduced carbon sink function of tropical forests. However, no study has established whether the liana effect on tree carbon accumulation is indeed stronger in the dry than in the wet season. Using 6 years of data from a large‐scale liana removal experiment in Panama, we provide the first experimental test of whether liana effects on tree carbon accumulation differ between seasons. We monitored tree and liana diameter increments at the beginning of the dry and wet season each year to assess seasonal differences in forest‐level carbon accumulation between removal and control plots. We found that median liana carbon accumulation was consistently higher in the dry (0.52 Mg C ha−1year−1) than the wet season (0.36 Mg C ha−1year−1) and significantly so in three of the years. Lianas reduced forest‐level median tree carbon accumulation more severely in the wet (1.45 Mg C ha−1year−1) than the dry (1.05 Mg C ha−1year−1) season in all years. However, the relative effect of lianas was similar between the seasons, with lianas reducing forest‐level tree carbon accumulation by 46.9% in the dry and 48.5% in the wet season.
- NSF-PAR ID:
- 10461500
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Journal of Ecology
- Volume:
- 107
- Issue:
- 4
- ISSN:
- 0022-0477
- Page Range / eLocation ID:
- p. 1890-1900
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract Synthesis . Competition for water plays a larger role in the interaction between lianas and trees than previously hypothesized, as demonstrated by simulations from a process‐based vegetation model. -
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Abstract Over the past decades, tropical forests have experienced both compositional and structural changes. In the Neotropics, researchers at multiple sites have observed significant increases in the abundance and biomass of lianas (i.e. woody vines) relative to trees. However, the role of dynamics at early life stages in contributing to increasing liana abundance remains unclear.
We took advantage of a unique dataset on seedling dynamics over 16 years in ~20 000 1‐m2plots in a tropical forest in Panama to examine temporal and spatial trends in liana and tree seedling abundance.
We found that the relative abundance of liana seedlings increased across the study period, from 0.18 in 2001 to 0.24 in 2017. However, increases in liana seedling relative abundance appear to have levelled off in more recent years. The observed increases in liana relative abundance appear to be the result of both higher survival and higher recruitment rates of liana seedlings compared to tree seedlings.
Increasing liana abundance in the seedling layer was not explained by annual variation in dry season length, total rainfall or the proportion of area occupied by canopy gaps. In addition, liana seedlings did not exhibit a demographic advantage (i.e. higher recruitment or survival) over tree seedlings in dry habitats.
Synthesis. Our results reveal that seedling communities experienced important compositional changes in the past, but liana seedling relative abundance may have stabilized in recent years. Longer‐term monitoring is needed to determine whether tropical forests will continue to experience compositional changes that may alter forest structure and ecosystem function. -
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Abstract Early successional tropical forests could mitigate climate change via rapid accumulation of atmospheric carbon. However, liana (woody vine) abundance and biomass has been increasing in many tropical forests over the past decades, which may slow the speed at which secondary forests accumulate biomass. Lianas decrease biomass accumulation in tropical forests, and may have a particularly strong effect on young forests by stalling tree growth. As forests mature, trees may outgrow or shed lianas, thus escaping some of the negative effects of lianas. Alternatively, lianas may have the strongest effect in older successional forests if the effect of lianas is commensurate with their density, which increases dramatically in the first decades of forest succession. We tested these two hypotheses using a landscape liana‐removal experiment in 30 forest stands that ranged from 10 to 35 yr old in Central Panama. We measured tree growth and biomass accumulation in the stands every year from 2014 to 2017. We found that the effect of liana removal on large trees (≥20‐cm diameter) decreased with forest age, supporting the hypothesis that lianas have the strongest negative effects on trees, and thus biomass uptake and carbon storage, in very young successional forests. Large trees accumulated more biomass in the absence of lianas in younger forests than in older forests (compared to controls) even after accounting for the effect of canopy completeness and crown illumination, implying that the detrimental effects of lianas go well beyond resource availability and crown health. There was no significant effect of lianas on small trees (1–20‐cm diameter), likely because lianas seek light and thus do not deploy their leaves on small trees that are trapped in the forest understory. Our results show that high liana density early in forest succession reduces forest biomass accumulation by negatively impacting large trees, thus decreasing the capacity of young secondary forests to mitigate climate change. Although the negative effects of lianas on forest biomass diminish as forests age, they do not disappear, and thus lianas are an important component of tropical forest carbon budgets throughout succession.
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Abstract Lianas reduce tree growth, reproduction, and survival in tropical forests. Liana competition can be particularly intense in isolated forest fragments, where liana densities are high, and thus, host tree infestation is common. Furthermore, lianas appear to grow particularly well during seasonal drought, when they may compete particularly intensely with trees. Few studies, however, have experimentally quantified the seasonal effects of liana competition on multiple tree species in tropical forests. We used a liana removal experiment in a forest fragment in southeastern Brazil to test whether the effects of lianas on tree growth vary with season and tree species identity. We conducted monthly diameter measurements using dendrometer bands on 88 individuals of five tree species for 24 months. We found that lianas had a stronger negative effect on some tree species during the wet season compared to the dry season. Furthermore, lianas significantly reduced the diameter growth of two tree species but had no effect on the other three tree species. The strong negative effect of lianas on some trees, particularly during the wet season, indicates that the effect of lianas on trees varies both seasonally and with tree species identity.
Abstract in Portuguese is available with online material.
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The water table lies approximately 12–14 m below the surface. The climate is humid temperate with a mean annual air temperature of 9.1 °C and annual precipitation of 1005 mm, 511 mm of which falls between May and September (1981–2010)56,57. The BCSE was established as a randomized complete block design in 2008 on preexisting farmland. Prior to BCSE establishment, the field was used for grain crop and alfalfa (Medicago sativa L.) production for several decades. Between 2003 and 2007, the field received a total of ~ 300 kg P ha−1 as manure, and the southern half, which contains one of four replicate plots, received an additional 206 kg P ha−1 as inorganic fertilizer. The experimental design consists of five randomized blocks each containing one replicate plot (28 by 40 m) of 10 cropping systems (treatments) (Supplementary Fig. S1; also see Sanford et al.58). Block 5 is not included in the present study. 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Between 2009 and 2016, soil samples (0–25 cm depth) were collected each autumn from all plots for determination of soil test P (STP) by the Bray-1 method61, using as an extractant a dilute hydrochloric acid and ammonium fluoride solution, as is recommended for neutral to slightly acidic soils. The measured STP concentration in mg P kg−1 was converted to kg P ha−1 based on soil sampling depth and soil bulk density (mean, 1.5 g cm−3). Sampling of water samples from lakes, streams and wells for P determination In addition to chemistry of soil and subsurface soil water in the BCSE, waters from lakes, streams, and residential water supply wells were also sampled during 2009–2016 for TDP analysis using Supor 450 membrane filters and the same analytical method as for soil water. These water bodies are within 15 km of the study site, within a landscape mosaic of row crops, grasslands, deciduous forest, and wetlands, with some residential development (Supplementary Fig. S2, Supplementary Table S2). Details of land use and cover change in the vicinity of KBS are given in Hamilton et al.48, and patterns in nutrient concentrations in local surface waters are further discussed in Hamilton62. Leaching estimates, modeled drainage, and data analysis Leaching was estimated at daily time steps and summarized as total leaching on a crop-year basis, defined from the date of planting or leaf emergence in a given year to the day prior to planting or emergence in the following year. TDP concentrations (mg L−1) of subsurface soil water were linearly interpolated between sampling dates during non-freezing periods (April–November) and over non-sampling periods (December–March) based on the preceding November and subsequent April samples. Daily rates of TDP leaching (kg ha−1) were calculated by multiplying concentration (mg L−1) by drainage rates (m3 ha−1 day−1) modeled by the Systems Approach for Land Use Sustainability (SALUS) model, a crop growth model that is well calibrated for KBS soil and environmental conditions. SALUS simulates yield and environmental outcomes in response to weather, soil, management (planting dates, plant population, irrigation, N fertilizer application, and tillage), and genetics63. The SALUS water balance sub-model simulates surface runoff, saturated and unsaturated water flow, drainage, root water uptake, and evapotranspiration during growing and non-growing seasons63. The SALUS model has been used in studies of evapotranspiration48,51,64 and nutrient leaching20,65,66,67 from KBS soils, and its predictions of growing-season evapotranspiration are consistent with independent measurements based on growing-season soil water drawdown53 and evapotranspiration measured by eddy covariance68. Phosphorus leaching was assumed insignificant on days when SALUS predicted no drainage. 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 (https://lter.kbs.msu.edu/datatables/7). 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 (https://data.sustainability.glbrc.org/protocols/156) species plant species that are rooted within the quadrat during the time of maximum biomass harvest. See protocol for more information, refer to link (http://lter.kbs.msu.edu/datatables/36) 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)more » « less
