John S. King’s research while affiliated with North Carolina State University and other places

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Publications (142)


Spatial distribution of freshwater wetlands over (a) the Southeastern (SE) United States (US), (b) the Mississippi Delta, and (c) the Everglades, overlaid by four representative CH4 flux observation sites (FLUXNET‐CH4 sites) used in this study and state boundaries. The wetland percentages at a spatial resolution of 0.0083° × 0.0083° were calculated from the National Wetlands Inventory (NWI).
Random forest (RF) model estimated (y‐axis) versus measured (x‐axis) monthly mean wetland CH4 fluxes for sites: (a) US‐DPW, (b) US‐LA2, (c) US‐MAC, and (d) US‐NC4. The black line is the 1:1 line, and the red line is the best‐fit line between the simulated and measured wetland CH4 fluxes. R² represents the coefficient of determination, while MAE denotes the mean absolute error (unit: nmol m⁻² s⁻¹). (e) Variable importance analysis in the RF model based on Gini importance values. The x‐axis lists explanatory variables ranked in descending order of importance: air temperature (Tair), water level (WTL), wind speed (WS), fall (Fall), incoming shortwave radiation (SW_IN), vapor pressure deficit (VPD), summer (Summer), precipitation (P), spring (Spring), and winter (Winter). The y‐axis shows the relative importance of these variables in estimating wetland methane fluxes. Partial dependence of wetland CH4 fluxes (y‐axis) on dominant predictors (x‐axis), (f) Tair, (g) WTL, and (h) Tair and WTL combined. The dots in (f–h) are eddy covariance tower measurements from the four FLUXNET‐CH4 sites.
(a) Climatological mean and (b) variability of subtropical freshwater wetland CH4 emissions. CH4 flux variability in (b) was calculated as one standard deviation of the monthly CH4 emissions across subtropical freshwater wetlands in the SE US during 1982–2010 at a high spatial resolution of 0.0083° × 0.0083° (∼1 km × 1 km). These estimates were generated using the RF regression model and based on the NWI freshwater wetland percentage data. (c) Interannual and (d) monthly variations of CH4 emissions from SE US subtropical freshwater wetlands. The red line in (c) represents the best‐fit linear regression line between the simulated CH4 emissions and the year. The bars in (d) indicate one standard deviation of monthly emissions during 1982–2010.
Linear regressions between the CMS Methane Flux product‐derived and the RF regression model‐estimated subtropical freshwater wetland CH4 fluxes on (a) monthly and (c) annual scales in 2010. The black line is the 1:1 line, while the red line is the best‐fit line. Boxplots of paired t‐test results comparing the CMS Methane Flux product‐derived and the RF regression model‐estimated CH4 fluxes at the (b) monthly and (d) annual scales in 2010. The red solid horizontal line, box, and whisker ends indicate the median, 25th and 75th percentiles, and the 10th and 90th percentiles, respectively. The data points outside the ranges are shown by the red plus (+) sign. (e) Spatial correlations between the CMS Methane Flux product‐derived and the RF regression model‐estimated wetland CH4 fluxes on a monthly scale in 2010. Significant correlations at the 5% significance level are stippled.
Conceptual model illustrating the effects of air temperature (Tair) and water level (WTL) on methane emissions in subtropical freshwater wetlands. Under the same WTL conditions, (left panel) lower temperatures (below the lower bound of the critical Tair range, which is 24°C) result in less wetland CH4 emissions compared to (right panel) higher temperatures. Within each panel, once WTL surpasses the WTL threshold (0.1 m in low‐temperature conditions and −0.7 m in high‐temperature conditions), wetland CH4 emissions increase (indicated by more and larger bubbles and red arrows) along the gradients of WTL.
Temperature and Water Levels Collectively Regulate Methane Emissions From Subtropical Freshwater Wetlands
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March 2025

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Curtis J. Richardson

Wetlands are the largest and most climate‐sensitive natural sources of methane. Accurately estimating wetland methane emissions involves reconciling inversion (“top‐down”) and process‐based (“bottom‐up”) models within the global methane budget. However, estimates from these two model types are inherently interdependent and often reveal substantial discrepancies. To enhance the reliability of both approaches, we need a comprehensive understanding of wetland methane emissions and an independent high‐resolution long‐term flux data set. Here, we employed a data‐driven random forest approach to identify key variables influencing methane emissions from subtropical freshwater wetlands in the Southeastern United States. The model‐estimated monthly mean methane fluxes fit well with measured methane fluxes (R² = 0.67) at four representative FLUXNET‐CH4 wetland sites across the region. Variable importance analysis highlighted the sensitivity of subtropical freshwater wetland methane emissions to variations in both temperature and water levels. High temperatures facilitate methanogenesis by enhancing microbial activities, while elevated water levels maintain anaerobic conditions necessary for methane production. Notably, the response of methane emissions to water level fluctuations is contingent on temperature conditions, and vice versa. Moreover, we constructed the first high‐spatial‐resolution (∼1 km × 1 km) and long‐term (1982–2010) gridded regional wetland methane flux product for the Southeastern United States, estimating annual methane emissions from subtropical freshwater wetlands in the region at 4.93 ± 0.11 Tg CH4 yr⁻¹ for 1982–2010. This new benchmark product holds promise for validating and parameterizing uncertain wetland methane emission processes in bottom‐up models and provides improved prior information for top‐down models.

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Hydrologic Perturbation Is a Key Driver of Tree Mortality in Bottomland Hardwood Wetland Forests of North Carolina, USA

December 2024

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36 Reads

Bottomland hardwood wetland forests along the Atlantic Coast of the United States have been changing over time; this change has been exceptionally apparent in the last two decades. Tree mortality is one of the most visually striking changes occurring in these coastal forests today. Using 2009–2019 tree mortality data from a bottomland hardwood forest monitored for long-term flux studies in North Carolina, we evaluated species composition and tree mortality trends and partitioned variance among hydrologic (e.g., sea level rise (SLR), groundwater table depth), biological (leaf area index (LAI)), and climatic (solar radiation and air temperature) variables affecting tree mortality. Results showed that the tree mortality rate rose from 1.64% in 2009 to 45.82% over 10 years. Tree mortality was primarily explained by a structural equation model (SEM) with R² estimates indicating the importance of hydrologic (R² = 0.65), biological (R² = 0.37), and climatic (R² = 0.10) variables. Prolonged inundation, SLR, and other stressors drove the early stages of ‘ghost forest’ formation in a formerly healthy forested wetland relatively far inland from the nearest coastline. This study contributes to a growing understanding of widespread coastal ecosystem transition as the continental margin adjusts to rising sea levels, which needs to be accounted for in ecosystem modeling frameworks.


Phenology Across Scales: An Intercontinental Analysis of Leaf-Out Dates in Temperate Deciduous Tree Communities

October 2024

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418 Reads

Aim: To quantify the intra-community variability of leaf-out (ICVLo) among dominant trees in temperate deciduous forests, assess its links with specific and phylogenetic diversity, identify its environmental drivers and deduce its ecological consequences with regard to radiation received and exposure to late frost. Location: Eastern North America (ENA) and Europe (EUR). Time Period: 2009-2022. Major Taxa Studied: Temperate deciduous forest trees. Methods: We developed an approach to quantify ICVLo through the analysis of RGB images taken from phenological cameras. We related ICVLo to species richness, phylogenetic diversity and environmental conditions. We quantified the intra-community variability of the amount of radiation received and of exposure to late frost. Results: Leaf-out occurred over a longer time interval in ENA than in EUR. The sensitivity of leaf-out to temperature was identical in both regions (−3.4 days per °C). The distributions of ICVLo were similar in EUR and ENA forests, despite the latter being more species-rich and phylogenetically diverse. In both regions, cooler conditions and an earlier occurrence of leaf-out resulted in higher ICVLo. ICVLo resulted in ca. 8% difference of radiation received from leaf-out to September among individual trees. Forest communities in ENA had shorter safety margins as regards the exposure to late frosts, and were actually more frequently exposed to late frosts. Main Conclusions: We conducted the first intercontinental analysis of the variability of leaf-out at the scale of tree communities. North American and European forests showed similar ICVLo, in spite of their differences in terms of species richness and phylogenetic diversity, highlighting the relevance of environmental controls on ICVLo. We quantified two ecological implications of ICVLo (difference in terms of radiation received and exposure to late frost), which should be explored in the context of ongoing climate change, which affects trees differently according to their phenological niche.


Active Remote Sensing Assessment of Biomass Productivity and Canopy Structure of Short-Rotation Coppice American Sycamore (Platanus occidentalis L.)

July 2024

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1 Citation

The short-rotation coppice (SRC) culture of trees provides a sustainable form of renewable biomass energy, while simultaneously sequestering carbon and contributing to the regional carbon feedstock balance. To understand the role of SRC in carbon feedstock balances, field inventories with selective destructive tree sampling are commonly used to estimate aboveground biomass (AGB) and canopy structure dynamics. However, these methods are resource intensive and spatially limited. To address these constraints, we examined the utility of publicly available airborne Light Detection and Ranging (LiDAR) data and easily accessible imagery from Unmanned Aerial Systems (UASs) to estimate the AGB and canopy structure of an American sycamore SRC in the piedmont region of North Carolina, USA. We compared LiDAR-derived AGB estimates to field estimates from 2015, and UAS-derived AGB estimates to field estimates from 2022 across four planting densities (10,000, 5000, 2500, and 1250 trees per hectare (tph)). The results showed significant effects of planting density treatments on LIDAR- and UAS-derived canopy metrics and significant relationships between these canopy metrics and AGB. In the 10,000 tph, the field-estimated AGB in 2015 (7.00 ± 1.56 Mg ha⁻¹) and LiDAR-derived AGB (7.19 ± 0.13 Mg ha⁻¹) were comparable. On the other hand, the UAS-derived AGB was overestimated in the 10,000 tph planting density and underestimated in the 1250 tph compared to the 2022 field-estimated AGB. This study demonstrates that the remote sensing-derived estimates are within an acceptable level of error for biomass estimation when compared to precise field estimates, thereby showing the potential for increasing the use of accessible remote-sensing technology to estimate AGB of SRC plantations.


Fig. A1. Box plots of stem biomass, root biomass, leaf area and root to shoot ratio of poplar plants -determined by destructive sampling at the end of 42-days experiment -for the different interacting environmental treatments. The left and right vertical panel represent the soil water availability treatments (D, 40% watered, left, W, 100% watered, right). Each figure panel is split according to CO 2 concentration (A, 400 ppm -left, E, 700 ppm, right) and color-coded according to VPD levels (L, low VPD = 1 kPa -red; H, VPD = 3 kPa -blue). The black dots represent the outliers detected according to 1.5 times IQR method. Note the logarithmic y-axis scale.
Fig. A2. Box plots depicting the rate of net assimilation (A net ), stomatal conductance (g s ), intrinsic water use efficiency (WUE i ) and transpiration rate (Tr) of young poplars under different interacting environmental treatments. The box plotted values represent medians of seven weekly measurements. The left and right vertical panel represent the soil water availability treatments (D, 40% watered, left, W, 100% watered, right). Each figure panel is split according to CO 2 concentration (A, 400 ppm -left, E, 700 ppm, right) and color-coded according to VPD levels (L, VPD = 1 kPa -red, H, VPD = 3 kPa -blue). The black dots represent the outliers detected according to 1.5 times IQR method. Note the logarithmic y-axis scale. VPD: maximum vapour pressure deficit.
Post hoc comparison of net assimilation (A net ), stomatal conductance (g s ), transpiration, and an intrinsic water use efficiency (WUE i ) based on seven gas exchange measurement campaigns conducted weekly during the period from DOY 210 to 245. Values sharing the same letters indicate no significant differences (p > 0.05).
Elevated CO 2 concentration alleviates the negative effect of vapour pressure deficit and soil drought on juvenile poplar growth

June 2024

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88 Reads

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1 Citation

Central European Forestry Journal

The growth performance of short-rotation woody coppice (SRWC) is strongly influenced by successful establishment in the initial months after planting. Future climates, expected to be warmer due to elevated atmospheric CO 2 (eCO 2), may bring about more frequent soil droughts alongside increased vapour pressure deficit (eVPD). Hence, this growth chamber experiment aimed to explore the interactive effects of eVPD, eCO 2 , and soil drought on growth and physiology traits of juvenile hybrid poplars under warmer climates. Our findings with juvenile hybrid poplar J-105 revealed that eVPD resulted in reductions in leaf area (-21%), root (-20%) and stem biomass (-9%), as well as in net assimilation (-15%), stomatal conductance (-26%), and transpiration (-13%). However, these decreases were relatively minor compared to the compensating effect of eCO 2 , which generally exerted a stronger influence than eVPD. While soil drought emerged as the primary growth-limiting factor in our study, elevated VPD is not expected to pose a significant additional threat to central European SRWC plantations of juvenile hybrid poplars under future conditions of ongoing climate change.



Energy availability and leaf area dominate control of ecosystem evapotranspiration in the southeastern U.S

April 2024

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116 Reads

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7 Citations

Agricultural and Forest Meteorology

Evapotranspiration (ET) links water, energy, and carbon balances, and its magnitude and patterns are changing due to climate and land use change in the southeastern U.S. Quantifying the environmental controls on ET is essential for developing reliable ecohydrological models for water resources management. Here, we synthesized eddy covariance data from 24 AmeriFlux sites distributed across the southeastern U.S., comprising 162 site-years of flux data representing six representative ecosystems including cropland vegetation mosaic (CVM), deciduous broadleaf forests (DBF), evergreen needle-leaf forests (ENF), grasslands (GRA), savannas (SAV), and wetlands (WET). Our objectives were to assess the daily, seasonal, and annual variability in ET and to develop practical predictive models for regional applications in ecosystem service analysis. We evaluated the response of ET to climatic and biotic forcings including potential evapotranspiration (PET), precipitation (P), and leaf area index (LAI), and compared the performance of these empirical ET models based and those developed using machine learning algorithms. Our results showed that the mean daily ET varied significantly, ranging from 1.36 mm d − 1 in GRA to 2.30 mm d − 1 in SAV, with a numerical order : GRA < DBF < ENF < WET < CVM < SAV. In this humid region, mean annual PET exceeded P in 16 out of the 24 flux sites. Using the Budyko framework, we showed that ENF had the highest evaporative efficiency (ET/P). PET and leaf area index (LAI) emerged as the most influential factors explaining ET variability. Artificial neural networks (ANN) and random forest (RF) models demonstrated superior capabilities in predicting monthly ET across sites over generalized additive modeling (GAM) and multiple linear regression (MLR) methods. The present study confirmed that the Southeast region is generally 'energy limited', implying that atmospheric demand along with vegetation information can be used to reliably estimate monthly and annual ET. Our study provides valuable insights into how ET of specific ecosystems is controlled by climatic and land surface drivers, enabling the development of reliable predictive models for regional extrapolation of flux measurements in water resource management in the humid southeastern U.S. region.



Phenology across scales: an intercontinental analysis of leaf-out dates in temperate deciduous tree communities

November 2023

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345 Reads

Aim. To quantify the intra-community variability of leaf-out (ICVLo) among dominant trees in temperate deciduous forests, assess its links with specific and phylogenetic diversity, identify its environmental drivers, and deduce its ecological consequences with regard to radiation received and exposure to late frost. Location. Eastern North America (ENA) and Europe (EUR). Time period. 2009-2022 Major taxa studied. Temperate deciduous forest trees. Methods. We developed an approach to quantify ICVLo through the analysis of RGB images taken from phenological cameras. We related ICVLo to species richness, phylogenetic diversity and environmental conditions. We quantified the intra-community variability of the amount of radiation received and of exposure to late frost. Results. Leaf-out occurred over a longer time interval in ENA than in EUR. The sensitivity of leaf-out to temperature was identical in both regions (-3.4 days per°C). The distributions of ICVLo were similar in EUR and ENA forests, despite the latter being more species-rich and phylogenetically diverse. In both regions, cooler conditions and an earlier occurrence of leaf-out resulted in higher ICVLo. ICVLo resulted in a ca. 8% difference of radiation absorption over spring among individual trees. Forest communities in ENA had shorter safety margins as regards the exposure to late frosts, and were actually more frequently exposed to late frosts. Main conclusions. We conducted the first intercontinental analysis of the variability of leaf-out at the scale of tree communities. North American and European forests showed similar ICVLo, in spite of their differences in terms of species richness and phylogenetic diversity, highlighting the relevance of environmental controls on ICVLo. We quantified two ecological implications of ICVLo


Merging flux-variance with surface renewal methods in the roughness sublayer and the atmospheric surface layer

November 2023

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204 Reads

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4 Citations

Agricultural and Forest Meteorology

Two micrometeorological methods utilizing high-frequency sampled air temperature were tested against eddy covariance (EC) sensible heat flux (H) measurements at three sites representing agricultural, agro-forestry, and forestry systems. The two methods cover conventional and newly proposed forms of the flux-variance (FV) and surface renewal (SR) schemes of differing complexities. The sites represent measurements in surface, roughness, and roughness to surface transitional layers. Regression analyzes against EC show that the most reliable FV and SR forms estimate H with slopes within ±10% from unity and coefficient of determination R2>0.9 across all the three sites. The best performance of both FV and SR was found at the agricultural site with measurements well within the surface layer, while the worst was found for the tall forest with measurements within the roughness sublayer where its thickness needed to be additionally estimated. The main variable driving H in FV is the temperature variance, whereas in SR, it is the geometry of ramp-like structures. Since these structures are also responsible for most of the temperature variance, a novel FV-SR approach emerging from combining the methods is proposed and evaluated against EC measurements and conventional FV and SR schemes. The proposed FV-SR approach requiring only a single fast response thermocouple is potentially independent of calibration and ameliorates some of the theoretical objections that arise when combining ramp statistics with similarity arguments. The combination of methods also provides new insights into the contribution of coherent structures to the temperature variance and its dependence on atmospheric stratification. Other potential utility of the new method is to include it in multi-tool assessments of surface energy fluxes, since a convergence or divergence of the results has a high diagnostic value.


Citations (75)


... This is because some plants can maintain high g s (e.g., Rasheed et al. 2015;Marchin et al. 2016) leading to important drops in Ψ beyond thresholds inducing embolism (Schönbeck et al. 2022). Finally, the few studies that have explored VPD effects on tree structure have found reduced growth and leaf area (Lendzion and Leuschner (2008); Orság et al. (2024). Yet, while higher stomatal sensitivity to VPD is usually found in more drought-tolerant species (Grossiord et al. 2020;Novick 2024), the acclimation potential between species is unknown. ...

Reference:

Acclimation to High Vapor Pressure Deficit in Warmer Air Can Reduce Tree Vulnerability to Drought‐Induced Mortality
Elevated CO 2 concentration alleviates the negative effect of vapour pressure deficit and soil drought on juvenile poplar growth

Central European Forestry Journal

... X i is the normalized predictor variables, and b i denotes their corresponding standardized regression coefficients. Separate MLR models were constructed for ET, ET c , ET i , ET s to evaluate their respective driving factors [54]. ...

Energy availability and leaf area dominate control of ecosystem evapotranspiration in the southeastern U.S
  • Citing Article
  • April 2024

Agricultural and Forest Meteorology

... The atmospheric surface layer is subdivided into an inertial sublayer and a roughness sublayer. While the inertial sublayer is characterized by a more homogeneous airflow, the roughness sublayer exhibits greater coherence which makes the surface renewal (SR) theory a more suitable approach for modeling heat transfer [14][15][16][17]. It explains why on bare soils the aerodynamic gradient method of the Monin-Obukhov similarity theory (MOST) was traditionally used [18]. ...

Merging flux-variance with surface renewal methods in the roughness sublayer and the atmospheric surface layer

Agricultural and Forest Meteorology

... Forested wetlands represent another poorly-understood terrestrial ecosystem. In this collection, Wells et al. (2023) explored the potential driving factors of interannual variability (IAV) of net ecosystem exchange (NEE) in forested wetlands using the Total Ecosystem (TECO) model based on long-term ecological data from three loblolly pine plantations and a bottomland hardwood forest of contrasting stand age in wetland areas of the lower coastal plain of North Carolina. Their findings suggested that anomaly correlation between IAV of NEE and ecological processes served as a useful tool for assessing the specific drivers of annual variability in ecosystem-level carbon exchange, and incorporating dynamic ecological responses could help improve the models. ...

Attributing interannual variability of net ecosystem exchange to modeled ecological processes in forested wetlands of contrasting stand age

Landscape Ecology

... Given the significant contribution of bubbling emissions to the overall greenhouse gas budget, understanding and accurately quantifying methane (CH₄) emissions from anoxic sediments is essential for advancing climate change research and developing effective environmental policies. Several methods are commonly employed to measure ebullition CH₄ emissions, including gas traps (Montes-Pérez et al. 2022), eddy covariance towers (McNicol et al. 2023), a range of gas sensors mounted on drones and aircraft (Lindgren et al. 2019;Hollenbeck et al. 2021;Dooley et al. 2024;Vélez et al. 2024), echosounders (Ostrovsky et al. 2008), and satellite-based imaging (Walter et al. 2008;Mathews et al. 2020). While these techniques are valuable for estimating gas fluxes, they often fall short in providing detailed information about the spatial distribution, number and size of gas bubbles. ...

Upscaling Wetland Methane Emissions From the FLUXNET‐CH4 Eddy Covariance Network (UpCH4 v1.0): Model Development, Network Assessment, and Budget Comparison

... For large-scale farms with a bigger animal population, the traditional techniques of raising cattle require the use of manpower for animal management and monitoring, which can be costly. For farms with 100-500 animals, it is not cost-effective to employ two competent workers to manage 20 animals, for instance (Ile et al., 2023). Additionally, this results in inadequate management of animal health and feed, which lowers yield and degrades animal health. ...

Toward the Adoption of New Farming Systems among Farmers: A Case study of Short Rotation Woody Crops in North Carolina

Journal of Extension

... By 2050, energy crops will surpass firewood as the primary source of biomass energy, contributing approximately 20% of the global energy supply, with wood crops playing a major role in this contribution [113]. Many agricultural countries and regions, such as the United States, promote shortrotation woody crops as they can serve as feedstocks for bioenergy, provide farmers with a stable income stream, and simultaneously avoid frequent soil disturbance while enhancing soil health, including improvements in soil structure, microbial lifespan, nutrient density, and carbon levels [114]. ...

Integrating Short Rotation Woody Crops into Conventional Agricultural Practices in the Southeastern United States: A Review

... Recent research has underscored the ecological significance of fine root turnover, particularly in forests experiencing seasonal droughts or nitrogen deposition [17]. However, variability in methodologies for studying fine root production, mortality, and decomposition calls for standardized approaches to improve comparability across studies [18]. Fine root decomposition has also been identified as a crucial link between soil organic matter dynamics and forest management practices, such as thinning intensity and recovery age. ...

Effects of methodological difference on fine root production, mortality and decomposition estimates differ between functional types in a planted loblolly pine forest

Plant and Soil

... This may be related to the difference in response mechanism between Ra and Rh to environmental factors. Rh is more sensitive to changes in environmental factors, especially in response to temperature changes (Wei et al. 2010;Liu et al. 2016Liu et al. , 2019aNoh et al. 2016;Kukumägi et al. 2017;Huang et al. 2021;Zheng et al. 2021;Morkoc et al. 2022). Temperature rise and precipitation increase within a certain threshold can significantly stimulate microbial activity, resulting in significant growth of Rh. ...

Environmental and Plant-Derived Controls on the Seasonality and Partitioning of Soil Respiration in an American Sycamore (Platanus occidentalis) Bioenergy Plantation Grown at Different Planting Densities

... WTL for US-LA2 were retrieved from the Coastwide Reference Monitoring System (co-located monitoring well CRMS2825-H01). WTL at US-MAC and US-NC4 were monitored on-site using ultrasonic water level dataloggers (Infinities, Port Orange, FL, USA; Aguilos et al., 2020Aguilos et al., , 2022. Monthly LAI and fAPAR were derived from the NOAA Climate Data Record (CDR) of Advanced Very High Resolution Radiometer (AVHRR) Surface Reflectance. ...

The Unabated Atmospheric Carbon Losses in a Drowning Wetland Forest of North Carolina: A Point of No Return?