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Tree radial growth is one of the most direct measures of tree performance and is also sensitive to climate. Growth performance is the consequence of the interplay between ecological and evolutionary processes. However, the effect of the evolutionary relatedness among species (i.e., phylogeny) on tree radial growth, especially under stressful conditions, remains largely unknown. Furthermore, there is still no ecological evidence for the influence of phylogeny on tree growth across different tree attributes (i.e., tree diameter variation and tree canopy height) and topographic habitat types. We used Blomberg's K to quantify the tree growth phylogenetic signal (TGPS) using two long‐term dendrometer data sets: one a continuous census of 225 tree species at 3‐month intervals in a tropical forest in southwest China from 2009 to 2017; the other, 12 tree species measured at 6‐month intervals in a temperate forest in Washington State, USA from 2013 to 2019. We found that TGPS values were higher in the temperate forest than in the tropical forest. Precipitation, tree diameter, canopy strata, and habitat types all influenced TGPS values. TGPS values were significantly (p < 0.05) and negatively related to precipitation in Xishuangbanna, and the three of four tree diameter classes in the temperate forest, respectively. Stressful growing conditions arose from either based on low precipitation or among large‐diameter trees competing with each other in the upper canopy led to phylogenetic conservatism in trees' radial growth performance. We conclude that phylogeny is pivotal to understanding the growth response differences among species and their responses to climate variability. Key Points: This study examined phylogeny‐based tree radial growth across precipitation gradients and tree attributes (i.e., tree diameter and canopy height) in tropical and temperate forestsTemperate trees showed a stronger phylogenetic growth effect than tropical treesTree radial growth phylogenetic conservatism was higher during stress events (i.e., drought) than in normal climatic situations [ABSTRACT FROM AUTHOR]

Invasive plants often alter ecosystem function and processes, especially soil N cycling. In eastern United States forests, the shrub Rosa multiflora ("rose") is a dominant invader, yet potential effects on N cycling are poorly understood. Moreover, invasive plant management can impact soil N cycling by decreasing plant N uptake and disturbing the soil. The objectives of this study were to evaluate N cycling along a gradient of rose invasion (observational) and investigate potential changes to N cycling (manipulative) under four different management strategies: (1) do nothing (the control), (2) invasive plant removal, (3) removal followed by native seed mix addition, (4) removal, native seed mix, and chipped rose stem addition. We selected three forest sites experiencing a Low, Medium, or High amount of shrub invasion, and measured N cycling in the early (June) and late (September) growing seasons. We found N was immobilized in June and mineralized in September. One year after experimental management, removal alone had no effect on N cycling compared to control plots, but addition of native seed mix and chipped stems reduced early-season nitrification in our Medium invasion site. Our findings suggest that rose invasion may increase N cycling rates when soils are dry, which may occur more frequently with future climate change. In addition, N cycling responds differentially to management in the year following invasive plant removal, but most noticeably under moderate rose invasion. [ABSTRACT FROM AUTHOR]

Invasive fungal pathogens are an increasing problem globally and can cause strong effects on forest ecosystems. In this study, we contrast vegetation surveys in eutrophic elm (Ulmus glabra) and ash (Fraxinus excelsior) forests in southern Sweden, conducted just prior to the arrival of Dutch elm disease (DED) in 1989, and then again in 2021, several years after ash dieback (ADB) began. At the sample plot scale, species richness (α–diversity) of the upper tree layer strongly decreased from 1989 to 2021, and the mean cover of elm decreased from 27 to 1% and of ash from 29 to 13%. In the lower tree and shrub layers, elm and ash were replaced by other, mainly shade-tolerant, tree species. The cover and richness of the shrub layer increased in previously elm-dominated stands but not in ash-dominated stands. The extensive loss of canopy cover in elm stands caused a larger change in upper tree layer species composition and increased compositional variability (β-diversity) between plots when compared to the ash stands. The direction of the changes in tree layer composition between the surveys varied with soil moisture and nutrient availability. While beech increased in less eutrophic plots, more nutrient-rich plots changed toward hornbeam or small-leaved lime, and wetter plots turned toward alder and bird cherry. Hence, our results indicate increased compositional diversity and alternative successional pathways for community reorganization following DED and ADB. Future research will reveal if these pathways will later merge or further split. [ABSTRACT FROM AUTHOR]

Global observations of solar‐induced chlorophyll fluorescence (SIF) are available from multiple satellite platforms, and SIF is increasingly used as a proxy for photosynthetic activity and ecosystem productivity. Because the relationship between SIF and gross primary productivity (GPP) depends on a variety of factors including ecosystem type and environmental conditions, it is necessary to study SIF observations across various spatiotemporal scales and ecosystems. To explore how SIF signals relate to productivity over a temperate deciduous forest, we deployed a PhotoSpec spectrometer system at the University of Michigan Biological Station AmeriFlux site (US‐UMB) in the northern Lower Peninsula of Michigan during the 2018 and 2019 growing seasons. We found that SIF correlated with GPP across diurnal and seasonal cycles (R2 = 0.61 and 0.64 for 90‐min‐ and daily‐averaged data), but that SIF signals were more strongly related to downwelling radiation than GPP (R2 = 0.91 for daily‐averaged data). The dependence of SIF on radiation obscured the impact of intraseasonal drought in the SIF timeseries, but drought stress was apparent as a decrease in relative SIF, which exhibited a stronger correlation with GPP (R2 = 0.56) than other remotely sensed data over the drought period. These results highlight the potential of SIF for detecting stress‐induced losses in forest productivity. Additionally, we found that the red:far‐red SIF ratio did not exhibit a response to water stress‐induced losses in productivity, but was largely driven by seasonal and interannual changes in canopy structure, as well as by synoptic changes in downwelling radiation. Plain Language Summary: Satellite measurements of solar‐induced chlorophyll fluorescence (SIF), a faint light signal emitted from vegetation during photosynthesis, are increasingly being used to estimate ecosystem productivity and carbon uptake. To accurately do so requires a robust understanding of how the relationship between SIF and plant productivity changes over time, in response to environmental stressors, and across different ecosystems. To better understand SIF signals and how they relate to carbon uptake over a temperate deciduous forest, we used a high‐precision spectrometer system to observe SIF signals at an AmeriFlux site (US‐UMB) in the northern Lower Peninsula of Michigan. While the shared dependence of SIF and ecosystem productivity on sunlight lead to strong daily and seasonal correlations, we found that SIF signals were more closely tied to the amount of incoming sunlight than to ecosystem productivity. Despite the stronger dependence of SIF on sunlight, we show that drought conditions lead to a lower SIF relative to the total light signal. Lastly, we show that the observation of SIF at multiple wavelengths may provide additional information on seasonal and interannual changes in canopy structure. Our results demonstrate the value and limitations in using SIF to assess carbon dynamics over temperate deciduous forest ecosystems. Key Points: Solar‐induced chlorophyll fluorescence above a temperate deciduous forest is more strongly tied to radiation than to productivityRelative solar‐induced fluorescence signals track water stress‐induced summer losses in productivity better than absolute fluorescenceThe ratio of red to far‐red solar‐induced fluorescence is sensitive to phenological changes in canopy structure and downwelling radiation [ABSTRACT FROM AUTHOR]

Trees continuously regulate leaf physiology to acquire CO2 while simultaneously avoiding excessive water loss. The balance between these two processes, or water use efficiency (WUE), is fundamentally important to understanding changes in carbon uptake and transpiration from the leaf to the globe under environmental change. While increasing atmospheric CO2 (iCO2) is known to increase tree intrinsic water use efficiency (iWUE), less clear are the additional impacts of climate and acidic air pollution and how they vary by tree species. Here, we couple annually resolved long‐term records of tree‐ring carbon isotope signatures with leaf physiological measurements of Quercus rubra (Quru) and Liriodendron tulipifera (Litu) at four study locations spanning nearly 100 km in the eastern United States to reconstruct historical iWUE, net photosynthesis (Anet), and stomatal conductance to water (gs) since 1940. We first show 16%–25% increases in tree iWUE since the mid‐20th century, primarily driven by iCO2, but also document the individual and interactive effects of nitrogen (NOx) and sulfur (SO2) air pollution overwhelming climate. We find evidence for Quru leaf gas exchange being less tightly regulated than Litu through an analysis of isotope‐derived leaf internal CO2 (Ci), particularly in wetter, recent years. Modeled estimates of seasonally integrated Anet and gs revealed a 43%–50% stimulation of Anet was responsible for increasing iWUE in both tree species throughout 79%–86% of the chronologies with reductions in gs attributable to the remaining 14%–21%, building upon a growing body of literature documenting stimulated Anet overwhelming reductions in gs as a primary mechanism of increasing iWUE of trees. Finally, our results underscore the importance of considering air pollution, which remains a major environmental issue in many areas of the world, alongside climate in the interpretation of leaf physiology derived from tree rings. [ABSTRACT FROM AUTHOR]

Interspecific interactions can mediate site occupancy of sympatric species and can be a key factor in habitat use patterns. American martens (Martes americana) and Fishers (Pekania pennanti) are two sympatric mesocarnivores in eastern North American forests. Due to their larger size, fishers have a competitive advantage over martens. We investigated site occupancy of martens and fishers in temperate deciduous forests of Québec, an environment modified by forest management and climate change. We formulated hypotheses on the spatial distribution of the studied species based on the knowledge of local trappers and on the scientific literature regarding forest cover composition, habitat fragmentation, and competitive relationships. We used a network of 49 camera traps monitored over two fall seasons to document site occupancy by both species. We used two-species site occupancy models to assess habitat use and the influence of fishers on martens at spatial grains of different sizes. None of the habitat variables that we considered explained site occupancy by fishers. Availability of dense old coniferous stands explained the spatial distribution of martens both at the home range grain size and at the landscape grain size. We identified the characteristics of habitat hotspots based on the knowledge of trappers, which highlighted the importance of stand composition, height, age, and canopy closure. The characteristics of habitat hotspots for martens in temperate deciduous forests refine the habitat suitability model for American martens that was originally developed for boreal forests of Québec. [ABSTRACT FROM AUTHOR]

We analyzed national forest inventory data collected from circa 2000-2018 across 37 states in the eastern United States to better understand the influence of forest health related canopy disturbances on the regeneration and recruitment dynamics of upland oaks (Quercus). We found low levels of oak recruitment across all disturbance types examined but limited evidence of any direct effects from the type of disturbance on the population of regenerating oaks. The general lack of differences in oak regeneration response between forest health disturbances and disturbances caused by harvested or non-disturbed plots does not indicate that the effects of forest health disturbances were benign, however. Instead, low level of oak recruitment across all disturbance types highlights the pervasiveness of the trend of shifting composition in once oak-dominated forests where oak is absent or sparse in the regeneration layer. Our results show that oak recruitment was higher when oak was present as advance reproduction prior to disturbance from any cause examined. Collectively, these results lead us to conclude that the widespread inadequacy of oak advance reproduction in mature oak-dominated forests is the prevailing threat to oak forest health and sustainability. We suggest the status of advance reproduction be treated as a co-morbidity when weighing the risk and potential outcomes from other threats to upland oak forests in the eastern United States. [ABSTRACT FROM AUTHOR]

Globally, temperate deciduous forests are threatened by invasion of non‐native (exotic) plant species. In the eastern United States, Rosa multiflora is a dominant shrub invader in forests, which often forms dense thickets that reduce sunlight availability in the understory, where decreased native plant diversity and abundance are observed. Management and restoration are difficult but desirable, especially when invasion intensity is still low. Few studies have examined the relative success of different management strategies under varying invasion intensities. Our study objectives were to conduct a R. multiflora removal experiment in three forest sites experiencing different invasion intensities and to restore native plant biodiversity while preventing secondary invasion. We utilized three management strategies: invasive plant removal, removal followed by native seed addition, and removal plus native seed and mulched invasive stem addition. We investigated the similarity between seed bank species composition and existing vegetation before and after removal to assess the potential for passive restoration. Two seasons after removal, we found that simply removing roses increased native species richness, native floristic quality assessment (FQAIN), and native shrub abundance in our medium invasion site, and total species richness in our low and medium invasion sites. Compared to removal alone, native seed addition, with and without mulch addition, resulted in larger native and total species richness and FQAIN increases at all sites, larger increases in native shrub abundance and exotic species richness in our medium invasion site, and larger reductions in exotic and total shrub abundance in our low and medium invasion sites. Following removal, species similarity between the seed bank and vegetation improved for all three sites. Our results indicate that removal of R. multiflora alone increased native plant biodiversity in the medium invasion scenario, but the seed bank may not provide a large native species pool. Additional management strategies lead to improved outcomes, especially in our most invaded forest, demonstrating the need to conduct multiple plant removal treatments across forests with varying site conditions and plant invasion intensity to improve management recommendations. [ABSTRACT FROM AUTHOR]

Key message: In mature Q. robur, chlorophyll varied with season and canopy height, whilst eCO2-driven changes were consistent with Marea, highlighting key factors for consideration when scaling photosynthetic processes and canopy N-use. Nitrogen-rich chlorophyll and carotenoid pigments are important in photosynthetic functioning. Photosynthetic pigments have been found to decrease with elevated CO2 (eCO2), but few such studies have been done in aged forest trees. This study aimed to assess the effects of eCO2 (150 μmol mol−1 above ambient) and canopy position on chlorophyll content in mature Quercus robur (Q. robur). Over 5000 in situ chlorophyll absorbance measurements, alongside laboratory chlorophyll extractions, were collected on canopy-dominant Q. robur in the 3rd and 4th season of CO2 fumigation of a free-air CO2 enrichment (FACE) study in central England. Mass-based chlorophyll concentration (Chlmass, mg g−1) was significantly higher in the lower canopy compared to upper canopy foliage (P < 0.05). In contrast, significantly higher chlorophyll content (Chlarea, mg m−2) was observed in the upper canopy. ECO2 did not affect Chlmass but Chlarea significantly increased, attributable to increased leaf mass per unit area (Marea, g m−2). We found no effect of eCO2 on mass-based or area-based nitrogen (Nmass, mg g−1 or Narea g m−2); however, Narea significantly increased with canopy height, again attributable to Marea. The parallel relationships between Marea, Narea and Chlarea suggest the allocation of N to light harvesting is maintained with eCO2 exposure as well as in the upper canopy, and that increased photosynthetic mass may help regulate the eCO2 variation. An understanding of changes in the light-harvesting machinery with eCO2 will be useful to assess canopy processes and, at larger scales, changes in biogeochemical cycles in future climate scenarios. [ABSTRACT FROM AUTHOR]

Abstract Globally, temperate deciduous forests are threatened by invasion of non‐native (exotic) plant species. In the eastern United States, Rosa multiflora is a dominant shrub invader in forests, which often forms dense thickets that reduce sunlight availability in the understory, where decreased native plant diversity and abundance are observed. Management and restoration are difficult but desirable, especially when invasion intensity is still low. Few studies have examined the relative success of different management strategies under varying invasion intensities. Our study objectives were to conduct a R. multiflora removal experiment in three forest sites experiencing different invasion intensities and to restore native plant biodiversity while preventing secondary invasion. We utilized three management strategies: invasive plant removal, removal followed by native seed addition, and removal plus native seed and mulched invasive stem addition. We investigated the similarity between seed bank species composition and existing vegetation before and after removal to assess the potential for passive restoration. Two seasons after removal, we found that simply removing roses increased native species richness, native floristic quality assessment (FQAIN), and native shrub abundance in our medium invasion site, and total species richness in our low and medium invasion sites. Compared to removal alone, native seed addition, with and without mulch addition, resulted in larger native and total species richness and FQAIN increases at all sites, larger increases in native shrub abundance and exotic species richness in our medium invasion site, and larger reductions in exotic and total shrub abundance in our low and medium invasion sites. Following removal, species similarity between the seed bank and vegetation improved for all three sites. Our results indicate that removal of R. multiflora alone increased native plant biodiversity in the medium invasion scenario, but the seed bank may not provide a large native species pool. Additional management strategies lead to improved outcomes, especially in our most invaded forest, demonstrating the need to conduct multiple plant removal treatments across forests with varying site conditions and plant invasion intensity to improve management recommendations.

We analyzed national forest inventory data collected from circa 2000–2018 across 37 states in the eastern United States to better understand the influence of forest health related canopy disturbances on the regeneration and recruitment dynamics of upland oaks (Quercus). We found low levels of oak recruitment across all disturbance types examined but limited evidence of any direct effects from the type of disturbance on the population of regenerating oaks. The general lack of differences in oak regeneration response between forest health disturbances and disturbances caused by harvested or non-disturbed plots does not indicate that the effects of forest health disturbances were benign, however. Instead, low level of oak recruitment across all disturbance types highlights the pervasiveness of the trend of shifting composition in once oak-dominated forests where oak is absent or sparse in the regeneration layer. Our results show that oak recruitment was higher when oak was present as advance reproduction prior to disturbance from any cause examined. Collectively, these results lead us to conclude that the widespread inadequacy of oak advance reproduction in mature oak-dominated forests is the prevailing threat to oak forest health and sustainability. We suggest the status of advance reproduction be treated as a co-morbidity when weighing the risk and potential outcomes from other threats to upland oak forests in the eastern United States.

This contribution discusses the optical properties of different structures of some herbaceous understorey plant species from temperate deciduous and mixed forests. These forests are marked by annual dynamics of radiation level that is related to the vegetation cycle of forest trees. During winter and early spring, the understorey is exposed to full solar radiation, whilelater in the growing season radiation is limited due to the closing of the tree storey. The plasticity of optical properties ofphotosynthetic structures of understorey plants is directly related to their structural and biochemical phenotypic plasticity that optimises harvesting and use of energy. The optimisation of energy harvesting is also achieved by specific adaptations of green leaves, such as variegation (Pulmonaria officinalis, Cyclamen sp.), anthocyanic lower epidermis (Cyclamen sp.), and by using structures other than green leaves for photosynthesis, such as bracts (Hacquetia epipactis) and sepals (Helleborus sp.). The optical properties of these structures are similar to those of green leaves. The understanding of optical responses of different structures contributes to the understanding of the forest understorey functioning.

Accurate individual tree segmentation (ITS) is fundamental to forest management and to the studies of forest ecosystem. Unmanned Aerial Vehicle Light Detection and Ranging (UAV-LiDAR) shows advantages for ITS and tree height estimation at stand and landscape scale. However, dense deciduous forests with tightly interlocked tree crowns challenge the performance for ITS. Available LiDAR points through tree crown and appropriate algorithm are expected to attack the problem. In this study, a new UAV-LiDAR dataset that fused leaf-off and leaf-on point cloud (FULD) was introduced to assess the synergetic benefits for ITS and tree height estimation by comparing different types of segmentation algorithms (i.e., watershed segmentation, point cloud segmentation and layer stacking segmentation) in the dense deciduous forests of Northeast China. Field validation was conducted in the four typical stands, including mixed broadleaved forest (MBF), Mongolian oak forest (MOF), mixed broadleaf-conifer forest (MBCF) and larch plantation forest (LPF). The results showed that the combination of FULD and the layer stacking segmentation (LSS) algorithm produced the highest accuracies across all forest types (F-score: 0.70 to 0.85). The FULD also showed a better performance on tree height estimation, with a root mean square error (RMSE) of 1.54 m at individual level. Compared with using the leaf-on dataset solely, the RMSE of tree height estimation was reduced by 0.22 to 0.27 m, and 12.3% more trees were correctly segmented by the FULD, which are mainly contributed by improved detection rate at nearly all DBH levels and by improved detection accuracy at low DBH levels. The improvements are attributed to abundant points from the bole to the treetop of FULD, as well as each layer point being included for segmentation by LSS algorithm. These findings provide useful insights to guide the application of FULD when more multi-temporal LiDAR data are available in future. [ABSTRACT FROM AUTHOR]

Context: Soil heterogeneity versus homogeneity patterns are observed within and across urban landscapes at multiple scales. To fully evaluate human-mediated influences on soil properties and processes, we need to understand spatial patterns and variation in soil characteristics within a single ecosystem patch type (e.g., forests) in and near cities. Objectives: Our research objectives were to: (1) identify soil characteristics important in driving variation in soil chemistry within urban forests, and (2) examine whether urbanization and invasion gradients were related to variation in soil chemistry within these forests. Methods: We measured soil chemical properties within 36 forests across the U.S. mid-Atlantic. The forests are spatially distributed across an urbanization gradient and comprise a gradient of non-native plant invasion. Results: Urbanization was related to more variation in soil chemistry, whereas plant invasion was related to less variation in soil chemistry within our forests. Soil Ca and Mg concentrations increase with plant invasion yet are less variable within invaded forests most likely due to invasive plants taking up and concentrating these elements. Soil pH, Ca, Mg, Zn, and Cu increase in forests surrounded by greater urbanization, however, these elements are more variable within urban forests likely due to edge effects altering element deposition. Conclusions: Our results demonstrate that while urbanization and invasion can increase soil chemical concentrations, they differentially alter variation in soil chemistry within urban forests. Plant invasion and urban environmental conditions need separate consideration in future conceptual models of urban ecological theory since non-native invasive plants influence soil chemistry independent of other urban factors. [ABSTRACT FROM AUTHOR]

Proper estimation of the herb layer annual net primary production (ANPP) can help to appreciate the role of this layer in carbon assimilation and nutrient cycling. Simple methods of ANPP estimation often understate its value. More accurate methods take into account biomass increments of individual species but are more laborious. We conducted our study in an oak-hornbeam forest (site area 12 ha) dominated by beech in NW Poland during two growing seasons (2010 and 2011). We collected herb layer biomass from 7 to 10 square frames (0.6 × 0.6 m). We collected plant biomass every week in April and May and every two weeks for the rest of the growing season. We compared six methods of calculating ANPP. The highest current-year standing biomass (1st method of ANPP calculation) was obtained on May 15, 2010—37.8 g m−2 and May 7, 2011—41.0 g m−2. The highest values of ANPP were obtained by the 6th method based on the sum of the highest products of shoot biomass and density for individual species: 74.3 g m−2 year−1 in 2010 and 94.0 g m−2 year−1 in 2011. The spring ephemeral Anemone nemorosa had the highest share of ANPP with 50% of the total ANPP. Two summer-greens, Galeobdolon luteum and Galium odoratum, each had a ca. 10% share of ANPP. The best results of ANPP calculation resulted from laborious tracking of dynamics of biomass and density of individual shoots. [ABSTRACT FROM AUTHOR]

In many agricultural landscapes, it is important to restore networks of forests to provide habitat and stepping stones for forest specialist taxa. More knowledge is, however, needed on how to facilitate the immigration of such taxa in restored forest patches. Here, we present the first chronosequence study to quantify the dynamics of immigration credits of forest specialist plants in post‐arable forest patches.We studied the distribution of herbaceous forest specialist plant species in 54 post‐arable broadleaved forest patches along gradients of age (20–140 years since forest establishment), distance from ancient forest (0–2,600 m) and patch area (0.5–9.6 ha). With linear mixed models, we estimated the effects of these factors on species richness, patch means of four dispersal‐related plant traits and with generalized linear models on the occurrence of 20 individual species.Post‐arable forest patch age and spatial isolation from ancient forest, but not patch size, were important predictors for species richness of forest specialists, suggesting that also small patches are valuable for habitat connectivity. Compared to species richness in ancient forest stands, the immigration credit was reduced by more than 90% after 80 years in post‐arable forest patches contiguous to ancient forest compared to 40% after 80 years and 60% after 140 years in isolated patches (at least 100 m to next forest). Tall‐growing species with adaptations to long‐distance dispersal were faster colonizers, whereas species with heavy diaspores and clonal growth were slower to colonize.Synthesis and applications. We show that post‐arable oak plantations have a high potential for restoration of forest herb vegetation. Dispersal‐related plant traits play a key role in explaining interspecific differences among forest specialists. To facilitate forest herb immigration across all functional groups in agricultural landscapes, we suggest to create clusters of relatively small new forest patches nearby older forest with source populations. [ABSTRACT FROM AUTHOR]

Previous studies found that pedunculate oak, one of the most widespread and abundant species in European deciduous forests, regenerates in open habitats and forest edges, but not in closed forest interiors. However, these observations usually come from the core areas of the biome, and much less is known about such processes at its arid boundary, where limiting factors may be different. In a full factorial field experiment, we tested the effects of different habitats (grassland, forest edge, forest interior) and increased growing season precipitation on the early regeneration of pedunculate oak in a forest-steppe ecosystem in Central Hungary, at the arid boundary of temperate deciduous forests. In the grassland habitat, seedling emergence was very low, and no seedlings survived by the fourth year. In contrast, seedling emergence was high and similar at forest edges and forest interiors, and was not affected by water addition. Most seedlings survived until the fourth year, with no difference between forest edge and forest interior habitats in numbers, and only minor or transient differences in size. The lack of oak regeneration in the grassland differs from previous reports on successful oak regeneration in open habitats, and may be related to a shift from light limitation to other limiting factors, such as moisture or microclimatic extremes, when moving away from the core of the deciduous forest biome towards its arid boundary. The similar number and performance of seedlings in forest edges and forest interiors may also be related to the decreasing importance of light limitation. [ABSTRACT FROM AUTHOR]

Accurate individual tree segmentation (ITS) is fundamental to forest management and to the studies of forest ecosystem. Unmanned Aerial Vehicle Light Detection and Ranging (UAV-LiDAR) shows advantages for ITS and tree height estimation at stand and landscape scale. However, dense deciduous forests with tightly interlocked tree crowns challenge the performance for ITS. Available LiDAR points through tree crown and appropriate algorithm are expected to attack the problem. In this study, a new UAV-LiDAR dataset that fused leaf-off and leaf-on point cloud (FULD) was introduced to assess the synergetic benefits for ITS and tree height estimation by comparing different types of segmentation algorithms (i.e., watershed segmentation, point cloud segmentation and layer stacking segmentation) in the dense deciduous forests of Northeast China. Field validation was conducted in the four typical stands, including mixed broadleaved forest (MBF), Mongolian oak forest (MOF), mixed broadleaf-conifer forest (MBCF) and larch plantation forest (LPF). The results showed that the combination of FULD and the layer stacking segmentation (LSS) algorithm produced the highest accuracies across all forest types (F-score: 0.70 to 0.85). The FULD also showed a better performance on tree height estimation, with a root mean square error (RMSE) of 1.54 m at individual level. Compared with using the leaf-on dataset solely, the RMSE of tree height estimation was reduced by 0.22 to 0.27 m, and 12.3% more trees were correctly segmented by the FULD, which are mainly contributed by improved detection rate at nearly all DBH levels and by improved detection accuracy at low DBH levels. The improvements are attributed to abundant points from the bole to the treetop of FULD, as well as each layer point being included for segmentation by LSS algorithm. These findings provide useful insights to guide the application of FULD when more multi-temporal LiDAR data are available in future.

Herbivore overabundance and species invasions could alter decomposition rates in temperate forests, with consequent effects on carbon sequestration, nutrient retention, and other ecosystem processes. At local scales, herbivores, invasive plants, and soil macroinvertebrates can be important drivers of decomposition, but interactive effects among these different groups are unknown. We tested for the effects of white-tailed deer, Amur honeysuckle, and earthworm activity (manipulated via mesh exclusion) on litter decomposition rates and loss of litter nitrogen (N) in five hardwood forest sites in southwestern Ohio. Each site consisted of a 20 × 20-m deer exclosure paired with a deer access plot; honeysuckle was removed from half of each plot. Effects of earthworm activity were tested using paired litter boxes of fine mesh (0.25 mm; earthworms excluded) or coarse mesh (10 mm; earthworm access). Restriction of earthworm activity in fine mesh treatments slowed litter decomposition and increased retention of N in the litter layer compared to coarse mesh. Deer access interacted with mesh treatments, with faster decomposition occurring in deer access, coarse mesh treatments relative to others. Greater earthworm biomass in deer access plots relative to deer exclosure plots corresponded with more rapid litter decomposition. Honeysuckle presence did not affect litter decomposition, but did increase litter N retention. The interactions between deer and earthworm activity indicate that reductions in deer populations may slow litter decomposition rates, increasing complexity of habitat structure at the soil surface, which relates to habitat for plants and animals. [ABSTRACT FROM AUTHOR]

Secondary forest regrowth shapes community succession and biogeochemistry for decades, including in the Upper Great Lakes region. Vegetation models encapsulate our understanding of forest function, and whether models can reproduce multi‐decadal succession patterns is an indication of our ability to predict forest responses to future change. We test the ability of a vegetation model to simulate C cycling and community composition during 100 years of forest regrowth following stand‐replacing disturbance, asking (a) Which processes and parameters are most important to accurately model Upper Midwest forest succession? (b) What is the relative importance of model structure versus parameter values to these predictions? We ran ensembles of the Ecosystem Demography model v2.2 with different representations of processes important to competition for light. We compared the magnitude of structural and parameter uncertainty and assessed which sub‐model–parameter combinations best reproduced observed C fluxes and community composition. On average, our simulations underestimated observed net primary productivity (NPP) and leaf area index (LAI) after 100 years and predicted complete dominance by a single plant functional type (PFT). Out of 4,000 simulations, only nine fell within the observed range of both NPP and LAI, but these predicted unrealistically complete dominance by either early hardwood or pine PFTs. A different set of seven simulations were ecologically plausible but under‐predicted observed NPP and LAI. Parameter uncertainty was large; NPP and LAI ranged from ~0% to >200% of their mean value, and any PFT could become dominant. The two parameters that contributed most to uncertainty in predicted NPP were plant–soil water conductance and growth respiration, both unobservable empirical coefficients. We conclude that (a) parameter uncertainty is more important than structural uncertainty, at least for ED‐2.2 in Upper Midwest forests and (b) simulating both productivity and plant community composition accurately without physically unrealistic parameters remains challenging for demographic vegetation models. [ABSTRACT FROM AUTHOR]

Temperate deciduous forests in the United States are located in the most densely populated states across the northern and mid-Atlantic east coast. Land development and associated human activities result in small forests that are susceptible to anthropogenic influences, such as urbanization and non-native plant invasion.The overall objective of this study was to assess spatial and temporal drivers of forest vegetation structure and diversity in small forests embedded across developed landscapes. We assessed woody plant composition across spatial gradients (i.e., urbanization and non-native plant invasion) and a temporal gradient (i.e., time since canopy closure) across 38 forests along the east coast of the United States in northern Delaware and southeastern Pennsylvania.Surprisingly, we found the invasion gradient was not related to the urban gradient across our forests. Across all forests, the canopy consisted of native species (98% of all trees), whereas the forest understory was most vulnerable to non-native plant invasion (65% of all woody plant stems). Greater native species richness in forest canopies and understories with increasing urbanization supports the conclusion that urban forests maintain native species and are not inherently degraded ecosystems. Non-native plant invasion has a strong influence on understory plant communities, and the duration of intact forest canopy had a strong negative correlation with non-native plant invasion suggesting intact forests can resist invasion.Synthesis. This is the first study to compare simultaneously the importance of invasion and urbanization in determining plant community composition in forests embedded across developed landscapes, and to discover that younger forests harbor more invasive plants and urban forests maintain native plants. [ABSTRACT FROM AUTHOR]

This paper reports the data collected in censuses of trees, litter fall and vegetation in a mountain riparian forest, upstream of the Watarase river, Gunma Prefecture in central Japan. A 1‐ha permanent plot (100 m × 100 m) subdivided into horizontal 10 m × 10 m grid cells was established in the forest in 2005. Tree census with woody species ≥15 cm girth at breast height was conducted every year from 2005. Twenty‐five litter traps were set in the permanent plot. The samples of litter fall were collected monthly, except in winter, between 2005 and 2008. And we sorted into leaves, branches, reproductive organs and other plant material, at species level as much as possible. The percentage cover of vascular plants was recorded at each vertical stratification (tree layer, subtree layer, shrub layer and herb layer) in 2008 and 2014 at the plot. These data are valuable for elucidating long‐term forest dynamics at mountain upstream areas experiencing frequent ground disturbances. The complete data set for this abstract published in the Data Paper section of the journal is available in electronic format in MetaCat in JaLTER at http://db.cger.nies.go.jp/JaLTER/metacat/metacat/ERDP-2020-15.1/jalter-en. [ABSTRACT FROM AUTHOR]

Aims: To develop forest vegetation classification at the level of alliances and associations across the Hyrcanian ecoregion, Northern Iran, and to explore the effects of main environmental and geographic gradients on their distribution. Location: Hyrcanian ecoregion, Northern Iran. Methods: A database of 1,597 vegetation plots of mostly 400 m2 in size with a total of 802 vascular plant taxa was established, covering the whole geographic range of the Hyrcanian forests at altitudes ranging from −22 to 2,850 m a. s. l. An expert system was developed for automatic classification of vegetation plots into alliances and associations. Detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA) were used to determine the most important environmental and geographic gradients affecting species composition. Results: Twenty‐one associations and seven alliances of these forests, belonging to five orders and four classes, were defined. Among them, eleven associations and five alliances were described as new syntaxa. Alnion glutinosae and Smilaco excelsae‐Alnion barbatae, both distributed in the lowland belt, include swamp and wet forests, respectively. Parrotio persicae‐Carpinion betuli and Alnion subcordatae are respectively mesic and wet forests of the submontane belt. Solano kieseritzkii‐Fagion orientalis is a mesic beech forest in the montane belt, and finally, Quercion macrantherae, an open oak forest, and Centaureo hyrcanicae‐Carpinion orientalis, a dry hornbeam forest, occur in the upper‐montane belt. DCA and CCA analyses showed that the distribution of these alliances and associations is mainly related to altitude and mean annual temperature. Conclusions: Based on our results and comparison between the Hyrcanian and European forests, we propose a new, comprehensive syntaxonomic scheme for the Hyrcanian forests, supported by a classification expert system. Unlike previous studies, we linked the classification system to that of EuroVegChecklist because, though this area is outside of Europe, its vegetation is very similar to that of the European temperate forest vegetation. [ABSTRACT FROM AUTHOR]

Among environmental factors, snow is one that is not considered enough in the Hyrcanian forests. There was a heavy snowfall in central Hyrcanian forests in 2016 November which caused serious damages to the forest trees. To consider these damages, 70 sample plots of 40x40 meter were chosen with systematic random method in parcels 609, 629 and 632 in Liresara Forest district in Mazandaran province. In each plot, all damaged and undamaged trees and the form and the properties of damages were considered. According to the results, 9% of trees had been damaged by snow. Amongst the damages, stem breakage and uprooting were the most frequent, whereas bending of stump was the least frequent. The most damages occurred to Alnus subcordata (with 22 percent of whole individuals) and Carpinus betulus, and Acer velutinum and Diospyros lotus were slightly damaged. Furthermore, A. subcordata, C. betulus, Parrotia persica and Quercus castaneifolia were more severely broken, while Buxus hyrcana was more uprooted and A. cappadocicum was more bent. The altitude class of lower than 200 meter had significantly the most damages; but slope classes did not show any significant differences. Concerning aspects, it was clarified that the most damages were on northern slopes. Altitude was significantly effective on extent of damage of A. subcordata and C. betulus, whereas Aspect was effective on C. betulus, A. cappadocicum and Tilia begonifolia. Young trees, grown after shelter cut of decades 1980 and 1990, whose diameter was 15-35 cm, were considerably vulnerable to snow. [ABSTRACT FROM AUTHOR]

Leaf litter decomposition rates (LDRs) of understory woody species vary substantially across species in temperate forest ecosystems. Using litter traits and LDR data for 78 shrub and liana species from a previous study, plus an additional litterbag experiment with varying mesh bag sizes for a subset of 17 species with rapid LDRs, we report that litter traits have nonlinear effects on LDR. In addition, we show that mesofauna, including nonnative earthworms, in general increase LDR and that the effects are greater for species of high LDR. Our results suggest that the acceleration of LDR by the co-invasion of plant species with high LDR and soil biota can promote nutrient cycling, potentially disrupting the stability of native forest ecosystems. [ABSTRACT FROM AUTHOR]

Amplicon-based analysis of 16S rRNA genes and transcripts was used to assess the effect of tree species composition on soil bacterial community structure and function in a temperate deciduous forest. Samples were collected from mono and mixed stands of Fagus sylvatica (beech), Carpinus betulus (hornbeam), Tilia sp. (lime), and Quercus sp. (oak) in spring, summer, and autumn. Soil bacterial community exhibited similar taxonomic composition at total (DNA-based) and potentially active community (RNA-based) level, with fewer taxa present at active community level. Members of Rhizobiales dominated at both total and active bacterial community level, followed by members of Acidobacteriales, Solibacterales, Rhodospirillales, and Xanthomonadales. Bacterial communities at total and active community level showed a significant positive correlation with tree species identity (mono stands) and to a lesser extent with tree species richness (mixed stands). Approximately 58 and 64% of indicator operational taxonomic units (OTUs) showed significant association with only one mono stand at total and active community level, respectively, indicating a strong impact of tree species on soil bacterial community composition. Soil C/N ratio, pH, and P content similarly exhibited a significant positive correlation with soil bacterial communities, which was attributed to direct and indirect effects of forest stands. Seasonality was the strongest driver of predicted metabolic functions related to C fixation and degradation, and N metabolism. Carbon and nitrogen metabolic processes were significantly abundant in spring, while C degradation gene abundances increased from summer to autumn, corresponding to increased litterfall and decomposition. The results revealed that in a spatially homogenous forest soil, tree species diversity and richness are dominant drivers of structure and composition in soil bacterial communities.

Ungulate browse and invasive plants exert pressure on plant communities and alter the physical and chemical properties of soils, but little is known about their effects on litter-dwelling arthropods. In particular, ants (Formicidae) are ubiquitous in temperate forests and are sensitive to changes in habitat structure and resources. As ants play many functional roles, changes to ant communities may lead to changes in ecosystem processes. We conducted a long-term experiment that controlled white-tailed deer (Odocoileus virginianus) access and presence of an invasive understory shrub in deciduous forests located in southwestern Ohio, USA from 2011 to 2017. Several leaf-litter ant community responses and litter biomass were measured in five paired deer access and exclosure plots, each with a split-plot removal of Amur honeysuckle (Lonicera maackii). Ant abundance and species richness increased with time in deer exclosures, but not in deer access plots. Honeysuckle removal reduced abundance and richness of ants. There were additive effects of deer and honeysuckle on ant richness, and interactive effects of deer and honeysuckle on ant abundance. Deer exclusion reduced variation in ant composition relative to access plots. There was little evidence that treatments directly influenced species diversity of ants. However, all ant measures were positively related to litter biomass, which was greater in deer exclosures relative to access plots. Our results indicate strong indirect effects of herbivores and honeysuckle on litter-dwelling ants, mediated through changes in litter biomass and likely vegetation structure, which may alter ant-mediated ecosystem processes. [ABSTRACT FROM AUTHOR]

Studying interactions between distantly related species is necessary to understand the complexity of food webs. Generalist predator interactions, such as intraguild predation (IGP) and competition, can alleviate predation pressure and weaken top–down control that predators have on lower trophic levels. Centipedes (Chilopoda) and carabid beetles (Coleoptera) are common deciduous forest floor generalist predators that may interact by competing for resources beneath rocks and logs on the forest floor, especially during dry periods when prey become confined to such microhabitats. We used laboratory and field studies to determine whether the carabid beetle, Pterostichus stygicus (Say), and the centipede, Scolopocryptops sexspinosus (Say) co-occur under artificial cover. Additionally, a laboratory mesocosm experiment was used to examine competitive interactions in intra- and interspecific trials. There was significant negative co-occurrence of beetles and centipedes beneath cover objects in the field and laboratory. Pairings of S. sexspinosus and P. stygicus within mesocosms resulted in high mortality of P. stygicus , and reciprocal but asymmetric IGP. Centipedes maintained weight within solitary, intra- and interspecific mesocosm treatments, however, beetles lost mass in all treatments. Scolopocryptops sexspinosus responded more favorably to intra- and interspecific competition than did P. stygicus. Analysis of the leaf litter mesofauna indicated that these predators consumed similar prey in laboratory mesocosms. Our results suggest that species with very different trophic morphology have the potential to compete for shared microhabitat and prey. [ABSTRACT FROM AUTHOR]

Summary: The climate sensitivity of forest ecosystem woody productivity (ANPPstem) influences carbon cycle responses to climate change. For the first time, we combined long‐term annual growth and forest census data of a diverse temperate broadleaf deciduous forest, seeking to resolve whether ANPPstem is primarily moisture‐ or energy‐limited and whether climate sensitivity has changed in recent decades characterised by more mesic conditions and elevated CO2.We analysed tree‐ring chronologies across 109 yr of monthly climatic variation (1901–2009) for 14 species representing 97% of ANPPstem in a 25.6 ha plot in northern Virginia, USA.Radial growth of most species and ecosystem‐level ANPPstem responded positively to cool, moist growing season conditions, but the same conditions in the previous May–July were associated with reduced growth. In recent decades (1980–2009), responses were more variable and, on average, weaker.Our results indicated that woody productivity is primarily limited by current growing season moisture, as opposed to temperature or sunlight, but additional complexity in climate sensitivity may reflect the use of stored carbohydrate reserves. Overall, while such forests currently display limited moisture sensitivity, their woody productivity is likely to decline under projected hotter and potentially drier growing season conditions. [ABSTRACT FROM AUTHOR]

Amplicon-based analysis of 16S rRNA genes and transcripts was used to assess the effect of tree species composition on soil bacterial community structure and function in a temperate deciduous forest. Samples were collected from mono and mixed stands of Fagus sylvatica (beech), Carpinus betulus (hornbeam), Tilia sp. (lime), and Quercus sp. (oak) in spring, summer, and autumn. Soil bacterial community exhibited similar taxonomic composition at total (DNA-based) and potentially active community (RNA-based) level, with fewer taxa present at active community level. Members of Rhizobiales dominated at both total and active bacterial community level, followed by members of Acidobacteriales, Solibacterales , Rhodospirillales , and Xanthomonadales. Bacterial communities at total and active community level showed a significant positive correlation with tree species identity (mono stands) and to a lesser extent with tree species richness (mixed stands). Approximately 58 and 64% of indicator operational taxonomic units (OTUs) showed significant association with only one mono stand at total and active community level, respectively, indicating a strong impact of tree species on soil bacterial community composition. Soil C/N ratio, pH, and P content similarly exhibited a significant positive correlation with soil bacterial communities, which was attributed to direct and indirect effects of forest stands. Seasonality was the strongest driver of predicted metabolic functions related to C fixation and degradation, and N metabolism. Carbon and nitrogen metabolic processes were significantly abundant in spring, while C degradation gene abundances increased from summer to autumn, corresponding to increased litterfall and decomposition. The results revealed that in a spatially homogenous forest soil, tree species diversity and richness are dominant drivers of structure and composition in soil bacterial communities. [ABSTRACT FROM AUTHOR]

Highlights • Mean and inter-annual variability in net ecosystem productivity (NEP) higher in a homogeneous, even-aged 130 years old forest compared to a heterogeneous forest. • Seasonal ecosystem processes more sensitive to abiotic environmental variables in structurally homogeneous forest compared to structurally heterogeneous forest. • Higher inter-annual variability of NEP in homogeneous forest mainly attributable to a time-dependent growth trend and fruit production. Abstract The effects of structural diversity on the carbon dioxide exchange (CO 2) of forests has become an important area of research for improving the predictability of future CO 2 budgets. We report the results of a paired eddy covariance tower study with 11 years of data on two forest sites of similar mean stand age, near-identical site conditions, and dominated by beech trees (Fagus sylvatica), but with a very different stand structure (incl. age, diameter distribution, stocks of dead wood and species composition) because of different management regimes. Here we address the question of how management and related structural diversity may affect CO 2 fluxes, and tested the hypothesis that more structurally diverse stands are less sensitive to variations in abiotic and biotic drivers. Higher annual net ecosystem productivity (NEP) was observed in the managed, even-aged, and homogenous forest (585 ± 57.8 g C m−2 yr−1), than in the unmanaged, uneven-aged, and structurally diverse forest (487 ± 144 g C m−2 yr−1). About two-third of the difference in NEP between the sites was contributed by a higher annual gross primary productivity (GPP, 1627 ± 164 vs 1558 ± 118 g C m−2 yr−1) and one-third by a lower annual ecosystem respiration (Reco, 1042 ± 60 vs 1071 ± 96 g C m−2 yr−1) in the homogenous forest. Spring (April – May) and summer (June – July) were the two main seasons contributing to the overall annual differences between the sites, also, the sensitivities of seasonal NEP and GPP to environmental variables were stronger in the homogenous forest during those periods. Inter-annual variation of NEP was higher in the homogenous forest (coefficient of variation (CV) = 25%) compared to the heterogeneous forest (CV = 12%). At annual time scale, the higher variability of NEP in the homogenous forest is attributed to biotic factors such as fruit production and a time-dependent growth trend, outweighing differences in environmental sensitivities. [ABSTRACT FROM AUTHOR]

Background Few studies of edge effects on wildlife objectively identify habitat edges or explore non-linear responses. In this paper, we build on ground beetle (Coleoptera: Carabidae) research that has begun to address these domains by using triangulation wombling to identify boundaries in beetle community structure and composition at the edges of forest patches with residential developments. We hypothesized that edges are characterized by boundaries in environmental variables that correspond to marked discontinuities in vegetation structure between maintained yards and forest. We expected environmental boundaries to be associated with beetle boundaries. Methods We collected beetles and measured environmental variables in 200 m by 200 m sampling grids centered at the edges of three forest patches, each with a rural, suburban, or urban context, in Charlotte, North Carolina, USA. We identified boundaries within each grid at two spatial scales and tested their significance and overlap using boundary statistics and overlap statistics, respectively. We complemented boundary delineation with k-means clustering. Results Boundaries in environmental variables, such as temperature, grass cover, and leaf litter depth, occurred at or near the edges of all three sites, in many cases at both scales. The beetle variables that exhibited the most pronounced boundary structure in relation to edges were total species evenness, generalist abundance, generalist richness, generalist evenness, and Agonum punctiforme abundance. Environmental and beetle boundaries also occurred within forest patches and residential developments, indicating substantial localized spatial variation on either side of edges. Boundaries in beetle and environmental variables that displayed boundary structure at edges significantly overlapped, as did boundaries on either side of edges. The comparison of boundaries and clusters revealed that boundaries formed parts of the borders of patches of similar beetle or environmental condition. Discussion We show that edge effects on ground beetle community structure and composition and environmental variation at the intersection of forest patches and residential developments can be described by boundaries and that these boundaries overlap in space. However, our results also highlight the complexity of edge effects in our system: environmental boundaries were located at or near edges whereas beetle boundaries related to edges could be spatially disjunct from them; boundaries incompletely delineated edges such that only parts of edges were well-described by sharp transitions in beetle and/or environmental variables; and the occurrence of boundaries related to edges was apparently influenced by individual property management practices, site-specific characteristics such as development geometry, and spatial scale.

Abstract: Shifts in rainfall patterns due to climate change are expected to increase drought stress and mortality in forests. Natural and anthropogenic fire regimes are also changing, highlighting the need to understand the interactive effects of fire and drought on tree ecophysiological response and growth. Using rainout shelters, we imposed summer drought on natural and planted populations of Quercus alba juveniles located in periodically burned and unburned sites in Shawnee National Forest, IL, USA. A subset of planted juveniles was clipped to simulate topkill. We measured predawn leaf water potential (Ψpd), leaf gas exchange (Amax, gmax) and relative growth rate (RGR) across treatments to test two hypotheses: (H1) Fire reduces juvenile drought stress by improving water relations through increased root‐to‐shoot ratios after topkill, or (H2) fire exacerbates juvenile drought stress by promoting a warmer, drier microclimate or amplifying drought‐induced nitrogen (N) limitation. Burned stands had higher vapour pressure deficits and 13% lower soil inorganic N availability than unburned stands. Rainout shelters reduced soil moisture (0–45 cm) by 10%–24% relative to ambient conditions. Consistent with H2, small, natural resprouts in burned stands experienced greater drought stress than unburned juveniles, with a 7% decrease in leaf nitrogen content (LNC), a 21%–29% reduction in Amax and gmax and a 41% reduction in RGR under drought. Drought effects on unburned juveniles, in contrast, were limited to a 5% reduction in LNC and a neutral to positive effect on leaf gas exchange and RGR. Large natural juveniles were largely unaffected by drought. Recent resprouts (i.e., clipped, planted juveniles) experienced less drought stress than unclipped juveniles, providing partial support for H1. Collectively, these results suggest that resprouting after fire can temporarily improve leaf water relations until root‐to‐shoot ratios re‐equilibrate. In contrast, fire can exacerbate drought‐driven declines in the growth of small juveniles by both promoting a warmer, drier microclimate and intensifying N limitation. Our results suggest that despite the high drought tolerance of Quercus spp., fire‐driven changes to local microclimate and resource conditions could limit tree recruitment under future scenarios of rainfall variability. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Declining near-infrared (NIR) surface reflectance between early and late summer, here termed greendown, is a common, yet poorly understood phenomena in remote sensing time series of temperate deciduous forests. As revealed by phenology analysis of Landsat satellite data, there are strong spatial patterns in the rate of greendown across temperate deciduous forest landscapes, and analyzing these patterns could help advance our understanding of surface reflectance drivers. Within an oak-hickory ( Quercus spp . – Carya spp.) forest landscape in western Maryland, USA, we tested how spatial patterns in greendown related to potential drivers at the landscape-, tree crown- and leaf-levels. We found that 50% of the spatial variability in greendown was explained by landscape variables, with greendown particularly higher in locations with higher maximum greenness, more southerly aspects, or locations with greater abundance of white oak ( Quercus alba ). The importance of species composition as a driver of greendown was supported at the tree crown level, where, relative to three other tree species, white oak exhibited the most consistent trend toward more vertical leaf angles later in the season. At the leaf level, NIR reflectance decreased in productive sites where %N increased, and δ 13 C decreased, through the season. However, among all sites, there were no consistent seasonal trends in foliar NIR reflectance, and we found no correlation among leaf-level NIR reflectance and satellite-observed greendown. Collectively, these results suggest that the spatial variability of greendown in this oak-hickory forest is most strongly controlled by an interaction of topographic and species compositional drivers operating at the landscape and tree crown levels. We found spatial analysis of greendown to be a useful approach to explore landscape-, tree crown-, and leaf-level controls on surface reflectance, and thereby help translate land surface phenology data into predictions of ecosystem structure and functioning. [ABSTRACT FROM AUTHOR]

Community-level shifts in the distributions of plant functional traits associated with environmental change are expected to influence ecosystem functioning. However, few studies have identified traits that both respond to environmental change and affect ecosystem properties, thus limiting potential to scale the effects of environmental change through the community level. We measured canopy and understory plant functional traits, characterizing the most abundant functional trait value (community-weighted mean; CWM) and the functional diversity (FD), across a soil resource gradient in fire-managed mixed-deciduous forests to determine how traits both respond to a disturbance-resource gradient and affect stocks of active and stable soil organic carbon (SOC) fractions. We expected that understory traits would respond mainly to fire and canopy traits would respond mainly to soil resources. We further hypothesized that fire and resource conditions affect SOC stocks (1) through mass-ratio, by influencing trait abundance; (2) through non-additive effects, by influencing the FD of plant communities; or (3) directly, through either combustion or environmental controls on SOC stocks. Understory traits responded to soil resource conditions and fire, whereas only canopy CWM leaf dry matter content (LDMC) varied with resource conditions; no canopy traits varied with fire. Among the response traits, canopy CWM LDMC and diversity in the maximum height of the understory were related to SOC stocks, suggesting they play dual roles as response and effect traits. SOC stocks were primarily associated with mass-ratio effects from canopy leaf traits and secondarily with non-additive effects from the canopy and understory. There were also strong, fraction-dependent patterns in SOC stocks with fire disturbance. Repeatedly burned forests characterized by resource conservative traits (i.e., high canopy CWM LDMC) had a higher relative proportion of active SOC, whereas unburned forests characterized by resource acquisitive traits (i.e., high canopy CWM leaf nitrogen content) had a higher relative proportion of stable SOC. Our results suggest that canopy community-aggregated leaf traits and diversity in understory size traits can act as both response and effect traits in disturbed forests. Predicting forest SOC stocks using a response–effect trait framework will thus require knowledge of both canopy and understory trait distributions, as well as disturbance history. [ABSTRACT FROM AUTHOR]