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Particle Shape, Size, and Composition from Satellites

The MISR Research Aerosol (RA) retrieval algorithm produces (A) aerosol amount (optical depth – AOD), (B) an aerosol size constraint (Angstrom Exponent) (C) fraction of non-spherical particles, and (D) particle light-absorption (SSA). Near the Camp fire source region the particles tend to be large (low Angstrom Exponent) and non-spherical particles, probably from burning in the town of Paradise. Aerosols are smaller and more spherical downwind, probably as the plume mixes with smoke from burning forest.

aerosol↗

Forest residue harvest optimization: spanning the bridge between plant biology and biorefinery performance

Forestry residues have immense potential as alternative feedstocks to petroleum, yet their inherent complexity remains a major challenge to widespread use. Pairing the temporal rhythms of plant biology with biorefinery performance is critical to industrial-scale biorefinery development. Here, we provide the first report of a techno-economic analysis (TEA) and life cycle assessment (LCA) for a model integrated reductive catalytic fractionation (RCF)–molten salt hydrolysis process for forestry residues varying in tree part, species, and phenophase. All forestry residues resulted in net-negative greenhouse gas (GHG) emissions vs. comparable petroleum feedstocks, with GHG emissions potentially reduced >4.0× through composition-based feedstock selection (e.g., harvesting American beech bark in spring vs. summer). Moreover, American beech twigs/branchlets and bark in leafed and emergence phenophases, respectively, had 7.9× lower predicted phenolic minimum selling prices (MSPs) vs. other feedstocks and MSPs within the current global phenolic market range. Hemicellulose content and RCF yield emerged as key parameters impacting GHG emissions and biorefinery revenue, identifying hardwood twigs/branchlets in the leafed phenophase as optimal biofeedstocks. Biorefinery expenses were dominated by purchased equipment, raw materials, and utility costs, highlighting essential areas for future study. Notably, RCF reactor pressures drove 85–90% of equipment costs, but sensitivity analysis revealed that decreasing the pressure 20% could reduce the phenol MSP 4-fold. Structural carbohydrate dynamics were also investigated using a two-step acid hydrolysis method to resolve tissue- and species-level patterns in biomass composition throughout the year to enable harvest optimization based on TEA/LCA findings. Ultimately, elucidating the impact of biofeedstock dynamics on biorefinery performance enables harvest optimization, informed engineering design, and progress towards an expanded bioeconomy.

Shapiro, Alison J. [University of Delaware, Newark↗

Signatures of aerosol-cloud interactions in GiOcean: a coupled global reanalysis with two-moment cloud microphysics

Aerosols influence the Earth's radiative balance through direct interactions with radiation and by affecting cloud properties. Anthropogenic aerosols have led to cooling during the industrial era through aerosol–cloud interactions (ACI), including aerosol effects on cloud microphysical properties and the subsequent adjustments. However, large uncertainties remain in Earth system models (ESMs) regarding the magnitude of this cooling. In part, ESMs substantially disagree on cloud properties, thermodynamics, the hydrological cycle, and general circulation. Reanalysis provides a useful avenue for exploring the impact of ACI on clouds and radiation because its atmosphere is forced to match realistic conditions through the assimilation of observations. Here, we explore the impact of ACI on clouds in the GiOcean reanalysis – the first to incorporate aerosol-cloud interactions. We contrast variables important for ACI between GiOcean and satellite observations and develop 2-dimensional lookup tables of ACI for both using a source-sink budget perspective to attribute the changes in cloud droplet number (Nd) and liquid water path (LWP) to aerosol and meteorology. A compositing analysis using lookup tables shows that GiOcean captures key aspects of aerosol–cloud–precipitation interactions, including (1) activation of aerosol into cloud droplets, (2) effective precipitation scavenging of Nd, (3) suppression of precipitation by high Nd in regions with heavy aerosol emissions. In contrast, satellite observations do not exhibit clear patterns for processes (2) and (3). Random Forest analysis shows that interannual variability in Nd and LWP over the Northern Hemisphere ocean in GiOcean is primarily driven by precipitation, consistent with satellite observations.

37 INORGANIC, ORGANIC, PHYSICAL, AND ANALYTICAL CH↗

Spatial transportability of geobotanical spectral signatures associated with soil geochemical anomalies

Consideration is given to a specific problem of spectral signature extension through examination and comparison of multitemporal spectral data signatures developed over a generalized, along-strike geochemical trend located within eastern deciduous hardwood forest. The spectral geobotanical signatures associated with soil Pb, Cu, and Zn anomalies were examined in south-central Virginia. Spectral discrimination between a training pair of anomalous and control sites within the Mineral district was 100 percent successful. It is concluded that spectral geobotanical signatures serve to identify regions of lowered biomass, lowered vigor, and/or differing species composition.

Bell, R.↗

Paraburkholderia madseniana sp. nov., a phenolic acid-degrading bacterium isolated from acidic forest soil

RP11 T was isolated from forest soil following enrichment with 4-hydroxybenzoic acid. Cells of RP11 T are aerobic, non-sporulating, exhibit swimming motility, and are rods (0.8 µm by 1.4 µm) that often occur as diplobacillus or in short chains (3–4 cells). Optimal growth on minimal media containing 4-hydroxybenzoic acid (µ=0.216 hr -1 ) occurred at 30 °C, pH 6.5 or 7.0 and 0% salinity. Comparative chemotaxonomic, genomic and phylogenetic analyses revealed the isolate was distinct from its closest relative type strains identified as Paraburkholderia aspalathi LMG 27731 T , Paraburkholderia fungorum LMG 16225 T and Paraburkholderia caffeinilytica CF1 T . Strain RP11 T is genetically distinct from P. aspalathi , its closest relative, in terms of 16S rRNA gene sequence similarity (98.7%), genomic average nucleotide identity (94%) and in silico DNA–DNA hybridization (56.7%±2.8). The composition of fatty acids and substrate utilization pattern differentiated strain RP11 T from its closest relatives, including growth on phthalic acid. Strain RP11 T encoded the greatest number of aromatic degradation genes of all eleven closely related type strains and uniquely encoded a phthalic acid dioxygenase and paralog of the 3-hydroxybenzoate 4-monooxygenase. The only ubiquinone detected in strain RP11 T was Q-8, and the major cellular fatty acids were C 16:0 , 3OH-C 16:0 , C 17:0 cyclo, C 19:0 cyclo ω8c, and summed feature 8 (C 18:1 ω7c/ω6c). On the basis of this polyphasic approach, it was determined that strain RP11 T represents a novel species from the genus Paraburkholderia for which the name Paraburkholderia madseniana sp. nov. is proposed. The type strain is RP11 T (=DSM 110123 T =LMG 31517 T ).

Microbiology↗

Analysis of hyper-spectral AVIRIS image data over a mixed-conifer forest in Maine

An introduction to some of the potential uses of hyperspectral data for ecosystem analysis is presented. The examples given are derived from a digital dataset acquired over a sub-boreal forest in central Maine in 1990 by the NASA-JPL Airborne Visible and Infrared Imaging Spectrometer (AVIRIS) instrument gathers data from 400 to 2500 nm in 224 channels at bandwidths of approximately 10 nm. As a preview to the uses of the hyperspectral data, several products from this dataset were extracted. They range from the traditional false color composite made from simulated Thematic Mapper bands and the well known normalized difference vegetation index to much more exotic products such as fractions of vegetation, soil and shade based on linear spectral mixing models and estimates of the leaf water content at the landscape level derived using spectrum-matching techniques. Our research and that of many others indicates that the hyperspectral datasets carry much important information which is only beginning to be understood. This analysis gives an initial indication of the utility of hyperspectral data. Much work still remains to be done in algorithm development and in understanding the physics behind the complex information signal carried in the hyperspectral datasets. This work must be carried out to provide the fullest science support for high spectral resolution data to be acquired by many of the instruments to be launched as part of the Earth Observing System program in the mid-1990's.

Lawrence, William T.↗

Soil Microbes Trade-Off Biogeochemical Cycling for Stress Tolerance Traits in Response to Year-Round Climate Change

Winter air temperatures are rising faster than summer air temperatures in high-latitude forests, increasing the frequency of soil freeze/thaw events in winter. To determine how climate warming and soil freeze/thaw cycles affect soil microbial communities and the ecosystem processes they drive, we leveraged the Climate Change across Seasons Experiment (CCASE) at the Hubbard Brook Experimental Forest in the northeastern United States, where replicate field plots receive one of three climate treatments: warming (+5°C above ambient in the growing season), warming in the growing season + winter freeze/thaw cycles (+5°C above ambient +4 freeze/thaw cycles during winter), and no treatment. Soil samples were taken from plots at six time points throughout the growing season and subjected to amplicon (rDNA) and metagenome sequencing. We found that soil fungal and bacterial community composition were affected by changes in soil temperature, where the taxonomic composition of microbial communities shifted more with the combination of growing-season warming and increased frequency of soil freeze/thaw cycles in winter than with warming alone. Warming increased the relative abundance of brown rot fungi and plant pathogens but decreased that of arbuscular mycorrhizal fungi, all of which recovered under combined growing-season warming and soil freeze/thaw cycles in winter. The abundance of animal parasites increased significantly under combined warming and freeze/thaw cycles. We also found that warming and soil freeze/thaw cycles suppressed bacterial taxa with the genetic potential for carbon (i.e., cellulose) decomposition and soil nitrogen cycling, such as N fixation and the final steps of denitrification. These new soil communities had higher genetic capacity for stress tolerance and lower genetic capacity to grow or reproduce, relative to the communities exposed to warming in the growing season alone. Our observations suggest that initial suppression of biogeochemical cycling with year-round climate change may be linked to the emergence of taxa that trade-off growth for stress tolerance traits.

54 ENVIRONMENTAL SCIENCES↗

Advancing Molecular Level Understanding of Aerosol Processes in the Amazon and Integration with Modeling (Final Report)

The Amazon forest is being converted to urban and agricultural uses through land clearing including large scale burning. It is also the dominant source of biogenic hydrocarbons globally, which can chemically transform in the atmosphere to form secondary organic aerosol (SOA). Most aerosols (solid or liquid particles suspended in air) are organic and formed through secondary chemical processes. This means that SOA chemical composition and their physical properties can impact cloud formation, the hydrologic cycle, and radiative balance in this region. The 2014 GoAmazon field campaign at the DOE/ARM facility at T3 afforded study of chemical transformations in the region downwind of Manaus. Local biogenic hydrocarbons emissions are high, and their chemical oxidation can be studied with varying degrees of influence by the urban plume. We collected aerosol filter samples and made time-resolved molecular level measurements by deploying a sequential filter sampler and a Semi-Volatile Thermal desorption Aerosol Gas Chromatograph (SV-TAG) during Jan-Mar 2014 (wet season) and Aug-Oct 2014 (dry season).

54 ENVIRONMENTAL SCIENCES↗

Point Cloud Based Mapping of Understory Shrub Fuel Distribution, Estimation of Fuel Consumption and Relationship to Pyrolysis Gas Emissions on Experimental Prescribed Burns

Forest fires spread via production and combustion of pyrolysis gases in the understory. The goal of the present paper is to understand the spatial location, distribution, and fraction (relative to the overstory) of understory plants, in this case, sparkleberry shrub, namely its degree of understory consumption upon burn, and to search for correlations between the degree of shrub consumption to the composition of emitted pyrolysis gases. Data were collected in situ at seven small experimental prescribed burns at Ft. Jackson, an army base in South Carolina, USA. Using airborne laser scanning (ALS) to map overstory tree crowns and terrestrial laser scanning (TLS) to characterize understory shrub fuel density, both pre- and postburn estimates of sparkleberry coverage were obtained. Sparkleberry clump polygons were manually digitized from a UAV-derived orthoimage of the understory and intersected with the TLS point cloud-derived rasters of pre- and postburn shrub fuel bulk density; these were compared in relation to overstory crown cover as well as to ground truth. Shrub fuel consumption was estimated from the digitized images; sparkleberry clump distributions were generally found to not correlate well to the overstory tree crowns, suggesting it is shade-tolerant. Moreover, no relationship was found between the magnitude of the fuel consumption and the chemical composition of pyrolysis gases, even though mixing ratios of 25 individual gases were measured.

54 ENVIRONMENTAL SCIENCES↗

Remote sensing of Earth terrain

Progress report on remote sensing of Earth terrain covering the period from Jan. to June 1993 is presented. Areas of research include: radiative transfer model for active and passive remote sensing of vegetation canopy; polarimetric thermal emission from rough ocean surfaces; polarimetric passive remote sensing of ocean wind vectors; polarimetric thermal emission from periodic water surfaces; layer model with tandom spheriodal scatterers for remote sensing of vegetation canopy; application of theoretical models to active and passive remote sensing of saline ice; radiative transfer theory for polarimetric remote sensing of pine forest; scattering of electromagnetic waves from a dense medium consisting of correlated mie scatterers with size distributions and applications to dry snow; variance of phase fluctuations of waves propagating through a random medium; polarimetric signatures of a canopy of dielectric cylinders based on first and second order vector radiative transfer theory; branching model for vegetation; polarimetric passive remote sensing of periodic surfaces; composite volume and surface scattering model; and radar image classification.

Kong, J. A.↗

Effects of warming and drying of soils on the ectomycorrhizal community of a mixed Pinus contorta/Picea engelmannii stand in Yellowstone Park

Restriction Fragment Length Polymorphisms (RFLPs) analyses were used to determine patterns of change in ectomycorrhizal community structure response to seasonal warming and drying of soils. Soil cores (42 total, 21 from cold and wet soil in early June, and 21 from dry, warm soil in late August) were collected from replicate blocks in a mixed-conifer forest stand in Yellowstone. Results indicated no significant differences in species richness (2.62 species/core, SE 0.2 in June; 3.25, SE 0.2 in August), however there was a significant effect on ectomycorrhizal infection (P<0.05), mean number of EM tips/core was significantly lower in June (185.8, SE 34) than in August (337 SE 78). Data indicated no difference in overall EM fungal species composition, however among system dominants, two species (Cortinarius 9 and Cortinarius 10) were more abundant in August than in June (P<0.02). The remaining dominant fungal species exhibited no differences in relative abundance. Results are discussed in relation to soil fertility and composition.

Cullings, Kenneth↗

Stability of tropical forest tree carbon-water relations in a rainfall exclusion treatment through shifts in effective water uptake depth

Increasing severity and frequency of drought is predicted for large portions of the terrestrial biosphere, with major impacts already documented in wet tropical forests. Using a 4-year rainfall exclusion experiment in the Daintree Rainforest in northeast Australia, we examined canopy tree responses to reduced precipitation and soil water availability by quantifying seasonal changes in plant hydraulic and carbon traits for 11 tree species between control and drought treatments. Even with reduced soil volumetric water content in the upper 1 m of soil in the drought treatment, we found no significant difference between treatments for predawn and midday leaf water potential, photosynthesis, stomatal conductance, foliar stable carbon isotope composition, leaf mass per area, turgor loss point, xylem vessel anatomy, or leaf and stem nonstructural carbohydrates. While empirical measurements of aboveground traits revealed homeostatic maintenance of plant water status and traits in response to reduced soil moisture, modeled belowground dynamics revealed that trees in the drought treatment shifted the depth from which water was acquired to deeper soil layers. Finally, these findings reveal that belowground acclimation of tree water uptake depth may buffer tropical rainforests from more severe droughts that may arise in future with climate change.

54 ENVIRONMENTAL SCIENCES↗

Whole-tree dormant season nitrogen pools for different species receiving combinations of fertilization and irrigation after one short rotation

Our understanding of the accretion and distribution of tree tissue nitrogen (N) pools across a variety of species and genotypes suitable for short-rotation woody crop (SRWC) production in response to water and N availability remains limited. We measured dormant-season, rotation-length, whole-tree N pools for five tree genotypes from four species (two eastern cottonwood, Populus deltoides Bartr., genotypes; American sycamore, Platanus occidentalis L.; American sweetgum, Liquidambar styraciflua L.; and loblolly pine, Pinus taeda L.) receiving irrigation (I), fertilization (F), their combination (IF), or no resources manipulation (C). Our results demonstrate that foliar nitrogen concentration [N] responded to fertilization but was constrained within genotype-specific ranges and varied temporally. Tree genotypes differed in their composite and component tissue N content (N C ), and these differences mostly reflected tissue mass (i.e., larger components and trees resulted in higher N C ). Resource amendments (I, F, IF) resulted in up to 3.8-fold increases in N C compared with C, which were most pronounced for sycamore and sweetgum with F and IF treatments, respectively. By the end of the rotation, forest stands accumulated 73 to 452 kg N ha –1 in tree tissues with 40 – 78 % distributed aboveground and 22 – 60 % distributed belowground. A critical difference between genotypes was that all hardwoods exhibited larger belowground N distributions than the evergreen conifer. Our results stress the importance of belowground N pools and highlight differences among genotypes. Furthermore, our study underscores valuable information about N pools across genotypes suitable for SRWC production, which can be leveraged to inform fertilization plans and devise sustainable nutrient management as production expands across marginal lands.

54 ENVIRONMENTAL SCIENCES↗

Improving the MODIS LAI Compositing Using Prior Time-Series Information

The Moderate Resolution Imaging Spectroradiometer (MODIS) long-term leaf area index (LAI) products have significantly contributed to global energy fluxes, climate change, and biogeochemistry research. However, the maximum fraction of photosynthetically active radiation absorbed by vegetation (Max-FPAR) compositing strategy of the Collection 6 (C6) products dictates that the main or backup algorithm is always triggered by observations of different quality, which indirectly causes the observed instability in the LAI time-series. Based on MODIS daily LAI retrievals, this study develops a prior knowledge time-series compositing algorithm (PKA) using a linear kernel driven (LKD) model. Our results show that the newly proposed PKA can significantly improve the LAI composites compared to the Max-FPAR strategy using ground-based observations for validation. We found that the PKA performs better than Max-FPAR in various aspects (different sites, seasons, and retrieval index (RI) ranges), with R2 increasing from 0.69 to 0.76 and root means square error (RMSE) decreasing from 1.01 to 0.84 compared to GBOV ground truth. The same improvement was shown for the ground truth LAIs measured at the Honghe and Hailun sites in northeastern China, with R2 increasing from 0.23 to 0.41 and RMSE decreasing from 1.27 to 1.25. In addition, three newly proposed temporal uncertainty metrics (time-series stability, TSS and time-series anomaly, TSA and reconstruction error metric, RE (the proximity to the main RT-based retrievals)) were applied to compare the stability of LAI time-series before and after PKA implementation. We found that the time series stability of PKA LAI was improved, the time series anomalies were reduced, and the retrieval rates of the main algorithm were also greatly enhanced compared to Max-FPAR LAI. A case intercomparison for Max-FPAR-MODIS, Max-FPAR-VIIRS (Visible Infrared Imager Radiometer Suite), and PKA-MODIS LAIs in the Amazon Forest region showed that the PKA is also effective in improving the LAI retrieval over large regions with few qualified observations due to poor atmospheric conditions (RE decreased from 2.37/2.35 (Max-FPAR-MODIS/Max-FPAR-VIIRS) to 2.25 (PKA-MODIS) and RI increased from 61.94%/59.62% to 66.88%). The same improvement was seen in the BELMANIP 2.1 sites for almost all biomes except deciduous broadleaf forest, where the RE decreased from 1.85/2.13 to 1.15 overall. We note that the PKA has the potential to be easily implemented in the operational algorithms of subsequent MODIS and MODIS-like LAI Collections.

MODIS↗

Invasive wild pig ( Sus scrofa ) diets on barrier islands in the southeastern United States

BACKGROUND: Biological invasions are a leading cause of reductions in global biodiversity. Islands are particularly sensitive to invasions, which often result in cascading impacts throughout island communities. Wild pigs (Sus scrofa) are globally invasive and pose threats to numerous taxa and ecosystems, particularly for islands where they have contributed to declines of many endemic species. However, the impacts of wild pig diet on the flora and fauna remain understudied in many island systems. RESULTS: We used DNA metabarcoding of wild pig fecal samples to quantify the seasonal diet composition of wild pigs on three barrier islands in the southeastern United States. Wild pigs exhibited a diverse diet dominated by plants, but also including marine and terrestrial animals. The diet composition of plants varied seasonally and between islands. Consumption of invertebrates also changed seasonally, with a shift to coastal invertebrates, particularly crabs, in spring and summer. Vertebrates were found in <10% of samples, but spanned broad taxa including amphibians, fish, mammals, and reptiles. Species consumed by wild pigs indicate that wild pigs use a variety of habitats within barrier islands for foraging, including maritime forests, saltmarshes, and beaches. CONCLUSIONS: An observed shift to beach foraging during sea turtle nesting season suggests wild pigs have potential to hinder nesting success on islands without established management programs. These findings provide insight into the diverse diets of wild pigs on barrier islands and highlight the need for removal of wild pigs from sensitive island ecosystems because of their potential impacts to native plant and animal communities. © 2024 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

60 APPLIED LIFE SCIENCES↗

Ectomycorrhizal and Dark Septate Fungal Associations of Pinyon Pine Are Differentially Affected by Experimental Drought and Warming

Changing climates can cause shifts in temperature and precipitation, resulting in warming and drought in some regions. Although each of these factors has been shown to detrimentally affect forest ecosystems worldwide, information on the impacts of the combined effects of warming and drought is lacking. Forest trees rely on mutualistic root-associated fungi that contribute significantly to plant health and protection against climate stresses. We used a six-year, ecosystem-scale temperature and precipitation manipulation experiment targeted to simulate the climate in 2100 in the Southwestern United States to quantify the effects of drought, warming and combined drought and warming on the root colonization (abundance), species composition and diversity of ectomycorrhizal fungi (EMF), and dark septate fungal endophytes in a widespread woodland tree, pinyon pine (Pinus edulis E.). Our results show that pinyon shoot growth after 6 years of these treatments was reduced more by drought than warming. The combined drought and warming treatment reduced the abundance and diversity of EMF more than either treatment alone. Individual ectomycorrhizal fungal taxa, including the drought tolerant Cenococcum geophilum, were present in all treatments but the combined drought and warming treatment. The combined drought and warming treatment also reduced the abundance of dark septate endophytes (DSE), but did not affect their diversity or species composition. The current year shoot growth of the trees correlated positively with ectomycorrhizal fungal diversity, highlighting the importance of diversity in mutualistic relationships to plant growth. Our results suggest that EMF may be more important than DSE to aboveground growth in P. edulis, but also more susceptible to the negative effects of combined climate stressors.

59 BASIC BIOLOGICAL SCIENCES↗

The usefullness of ERTS-1 and supporting aircraft data for monitoring plant development in rangeland environments

The author has identified the following significant results. Preliminary analysis of ERTS-1 MSS imagery of annual and perennial rangeland in California yields the following observations: (1) Sufficient geomorphological detail can be resolved to differentiate upland and bottomland range sites in the foothill range areas. (2) Dry and green meadowland can be differentiated on MSS band 5. (3) Color composites prepared by NASA-Goddard were useful for locating perennial rangeland with varying amounts of herbaceous ground cover. (4) The ERTS-1 images received and interpreted cover nearly 50% of the state of California and show nearly two-thirds of the annual grassland type. (5) Satellite imagery obtained during the late summer season should be optimum for differentiating grassland from brushland and forested land. (6) The ERTS-1 imagery clearly shows areas which at one time were part of the annual grassland but which are now used for dry land farming (cropping of cereal grains). Similarly, the imagery show areas which have been converted from brushland to grassland.

Carneggie, D. M.↗

Context matters: Natural tree mortality can lead to neighbor growth release or suppression

Where competition suppresses tree growth, mortality of adjacent trees can release the surviving individuals, leading to a growth increase. However, primary forests are complicated systems, where trees interact in both competitive and facilitative ways mediated by their size, species, and the broad ecological context in which they grow. Thus, the magnitude and even direction of growth responses to the mortality of nearby trees may vary, which has implications for our understanding of community- and ecosystem-level dynamics following mortality events. Unfortunately, although many studies focus on the impacts of light availability and general crowding on tree growth, we know relatively little about the effects of naturally occurring mortality events on the growth of neighboring trees. To address this issue, we used 40 years of data from 15 permanent forest-monitoring plots in Mt Rainier old-growth forests, comparing observed to expected radial growth of individual trees following the death of their nearest neighbor. Although we found evidence of a general growth-release response, this was not universal among all trees, with small trees in particular exhibiting growth suppression (rather than release) following neighboring tree mortality. In addition to small size, growth-suppression was more likely if the dead neighbor was the same species, consistent with facilitative effects as mediated through belowground networks. At the stand level, the average growth release after nearest neighbor mortality was greatest in low-density stands with large trees, with elevation and community composition also playing a role. Decades more monitoring could reveal how long growth release (or suppression) is sustained by individual trees following neighboring mortality events, as well as potential response lags and the role of species identity in determining whether interactions with neighbors are competitive or facilitative. Nonetheless, our results suggest that although mature trees have competitive effects on their larger neighbors, they also have an important role in supporting the ingrowth of small trees. More broadly, we demonstrate that the nature of interactions between individual neighboring trees is highly context dependent.

54 ENVIRONMENTAL SCIENCES↗