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Lidar‐derived forest structural diversity (FSD) metrics—including measures of forest canopy height, vegetation arrangement, canopy cover (CC), structural complexity and leaf area and density—are increasingly used to describe forest structural characteristics and can be used to infer many ecosystem functions. Despite broad adoption, the importance of spatial resolution (grain and extent) over which these structural metrics are calculated remains largely unconsidered. Often researchers will quantify FSD at the spatial grain size of the process of interest without considering the scale dependency or statistical behaviour of the FSD metric employed.We investigated the appropriate scale of inference for eight lidar‐derived spatial metrics—CC, canopy relief ratio, foliar height diversity, leaf area index, mean and median canopy height, mean outer canopy height, and rugosity (RT)‐‐representing five FSD categories—canopy arrangement, CC, canopy height, leaf area and density, and canopy complexity. Optimal scale was determined using the representative elementary area (REA) concept whereby the REA is the smallest grain size representative of the extent. Structural metrics were calculated at increasing canopy spatial grain (from 5 to 1000 m) from aerial lidar data collected at nine different forested ecosystems including sub‐boreal, broadleaf temperate, needleleaf temperate, dry tropical, woodland and savanna systems, all sites are part of the National Ecological Observatory Network within the conterminous United States. To identify the REA of each FSD metric, we used changepoint analysis via segmented or piecewise regression which identifies significant changepoints for both the magnitude and variance of each metric.We find that using a spatial grain size between 25 and 75 m sufficiently captures the REA of CC, canopy arrangement, canopy leaf area and canopy complexity metrics across multiple forest types and a grain size of 30–150 m captures the REA of canopy height metrics. However, differences were evident among forest types with higher REA necessary to characterize CC in evergreen needleleaf forests, and canopy height in deciduous broadleaved forests.These findings indicate the appropriate range of spatial grain sizes from which inferences can be drawn from this set of FSD metrics, informing the use of lidar‐derived structural metrics for research and management applications. [ABSTRACT FROM AUTHOR]

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Many secondary deciduous forests of eastern North America are approaching a transition in which mature early‐successional trees are declining, resulting in an uncertain future for this century‐long carbon (C) sink. We initiated the Forest Accelerated Succession Experiment (FASET) at the University of Michigan Biological Station to examine the patterns and mechanisms underlying forest C cycling following the stem girdling‐induced mortality of >6,700 early‐successional Populus spp. (aspen) and Betula papyrifera (paper birch). Meteorological flux tower‐based C cycling observations from the 33‐ha treatment forest have been paired with those from a nearby unmanipulated forest since 2008. Following over a decade of observations, we revisit our core hypothesis: that net ecosystem production (NEP) would increase following the transition to mid‐late‐successional species dominance due to increased canopy structural complexity. Supporting our hypothesis, NEP was stable, briefly declined, and then increased relative to the control in the decade following disturbance; however, increasing NEP was not associated with rising structural complexity but rather with a rapid 1‐yr recovery of total leaf area index as mid‐late‐successional Acer, Quercus, and Pinus assumed canopy dominance. The transition to mid‐late‐successional species dominance improved carbon‐use efficiency (CUE = NEP/gross primary production) as ecosystem respiration declined. Similar soil respiration rates in control and treatment forests, along with species differences in leaf physiology and the rising relative growth rates of mid‐late‐successional species in the treatment forest, suggest changes in aboveground plant respiration and growth were primarily responsible for increases in NEP. We conclude that deciduous forests transitioning from early to middle succession are capable of sustained or increased NEP, even when experiencing extensive tree mortality. This adds to mounting evidence that aging deciduous forests in the region will function as C sinks for decades to come. [ABSTRACT FROM AUTHOR]

The pre-disturbance vegetation characteristics that predict carbon (C) cycling responses to disturbance are not well known. To address this gap, we initiated the Forest Resilience Threshold Experiment, a manipulative study in which more than 3600 trees were stem girdled to achieve replicated factorial combinations of four levels (control, 45, 65, and 85% gross defoliation) of disturbance severity and two disturbance types (targeting upper or lower canopy strata). Applying a standardized stability framework in which initial C cycling resistance to disturbance was calculated as the first-year natural log response ratio of disturbance and control treatments, we investigated to what extent pre-disturbance levels of species diversity, aboveground woody biomass, leaf area index, and canopy rugosity—a measure of structural complexity—predict the initial responses of subcanopy light-saturated leaf CO2 assimilation (Asat), aboveground wood NPP (ANPPw), and soil respiration (Rs) to phloem-disrupting disturbance. In the year following stem girdling, we found that above-ground C cycling processes, Asat and ANPPw, were highly resistant to increases in disturbance severity, while Rs resistance declined as severity increased. Disturbance type had no effect on first-year resistance. Pre-disturbance aboveground woody biomass, and canopy rugosity were positive predictors of ANPPw resistance and, conversely, negatively related to Rs resistance. Subcanopy Asat resistance was not related to pre-disturbance vegetation characteristics. Stability of C uptake processes along with Rs declines suggest the net C sink was sustained in the initial months following disturbance. We conclude that biomass and complexity are significant, but not universal, predictors of initial C cycling resistance to disturbance. Moreover, our findings highlight the utility of standardized stability measures when comparing functional responses to disturbance. [ABSTRACT FROM AUTHOR]

Aim: Canopy structural complexity, which describes the degree of heterogeneity in vegetation density, is strongly tied to a number of ecosystem functions, but the community and structural characteristics that give rise to variation in complexity at site to subcontinental scales are poorly defined. We investigated how woody plant taxonomic and phylogenetic diversity, maximum canopy height, and leaf area index (LAI) relate to canopy rugosity, a measure of canopy structural complexity that is correlated with primary production, light capture, and resource‐use efficiency. Location: Our analysis used 122 plots distributed across 10 ecologically and climatically variable forests spanning a > 1,500 km latitudinal gradient within the National Ecological Observatory Network (NEON) of the USA. Time period: 2016–2018. Taxa studied: Woody plants. Methods: We used univariate and multivariate modelling to examine relationships between canopy rugosity, and community and structural characteristics hypothesized to drive site and subcontinental variation in complexity. Results: Spatial variation in canopy rugosity within sites and across the subcontinent was strongly and positively related to maximum canopy height (r2 =.87 subcontinent‐wide), with the addition of species richness in a multivariate model resolving another 2% of the variation across the subcontinent. Individually, woody plant species richness and phylogenetic diversity (r2 =.17 to.44, respectively) and LAI (r2 =.16) were weakly to moderately correlated with canopy rugosity at the subcontinental scale, and inconsistently explained spatial variation in canopy rugosity within sites. Main conclusions: We conclude that maximum canopy height is a substantially stronger predictor of complexity than diversity or LAI within and across forests of eastern North America, suggesting that canopy volume places a primary constraint on the development of structural complexity. Management and land‐use practices that encourage and sustain tall temperate forest canopies may support greater complexity and associated increases in ecosystem functioning. [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]

• Structural and composition change following stem-girdling varied by disturbance severity. • Removal of large trees affected soil temperature, soil water content, and canopy light absorption. • Removal of small to medium trees primarily affected only soil temperature. Forest structural diversity and community composition are key in regulating forest microclimates. When disturbance affects structural diversity or composition, forest microclimates may be altered due to changes in soil temperature, soil water content, and light availability. It is unclear however which structural or compositional components, when changed or to what extent, result in microclimatic change. To address this question, we used data from a large scale, manipulative stem-girdling experiment in northern, lower Michigan—the Forest Resilience and Threshold Experiment (FoRTE). FoRTE follows a factorial design with multiple levels of disturbance severity (0, 45, 65, 85%) based on targeted reductions in gross leaf area index via stem-girdling induced mortality. These disturbance severity treatments are applied in two ways: either as top-down (largest trees are killed) or bottom-up (small to medium trees killed) treatments. We examined how multiple components of structural diversity and community composition changed as a product of disturbance severity and type, and then tested for resulting effects on forest microclimates (light availability, soil temperature, and soil water), using a multivariate, Random Forest framework. We found that measures of community composition (species richness, species evenness, and Shannon-Wiener Diversity Index) and stand structure (basal area, standard deviation of DBH, tree size diversity) declined more following disturbance than did measures of canopy cover, heterogeneity, arrangement, or height. However, when changes in each variable from pre- to post-disturbance, measured as log change, were employed in a multivariate, Random Forest regression framework, structural diversity measures of heterogeneity (rugosity, top rugosity), cover (canopy cover), and arrangement (porosity) were the most influential variables, but with differences among bottom-up and top-down treatments We found that the death of large trees from disturbance impacts soil temperature, water, and light environments more substantially and uniformly across disturbance gradients than does the death of smaller trees. Our results have implications for both statistical and process-based modeling of forest disturbance. [ABSTRACT FROM AUTHOR]

• Disturbance effects on forest canopy structural complexity are poorly understood. • We synthesized the results of three forest disturbance manipulations. • Disturbance had variable effects on the temporal dynamics of structural complexity. • Moderate severity disturbance sometimes increased structural complexity. • Disturbance severity, source, and scale constrain temporal changes in complexity. The temporal dynamics of forest canopy structure are influenced by disturbances that alter vegetation quantity and distribution. While canopy structural indicators such as leaf area index (LAI), canopy cover, and canopy height have been widely studied in the context of disturbance, the post-disturbance temporal dynamics of structural complexity, which summarizes the heterogeneity of vegetation arrangement, are poorly understood. With the goal of advancing conceptual and empirical understanding of the temporal dynamics of structural complexity following disturbance, we synthesized results from three large-scale disturbance manipulation experiments at the University of Michigan Biological Station (UMBS): the 4-year Forest Resilience Threshold Experiment (FoRTE) manipulating levels of disturbance severity; the decade-long Forest Accelerated Succession Experiment (FASET), in which all early successional tree species were stem-girdled within 39 ha in the same landscape; and forest chronosequences established following clear-cut harvesting. We found that the temporal dynamics of canopy structure following disturbance were dependent upon three factors: (1) the source and severity of disturbance; (2) the spatial and temporal scales of analysis; and (3) the measure of structure assessed. Unlike vegetation area index and canopy cover, which initially decreased in response to disturbance, structural complexity measures such as canopy and top rugosity did not consistently respond to moderate levels of disturbance severity. Over multi-decadal timescales, structural complexity increased to a maximum, regardless of whether fire occurred at the time of stand establishment, but intervening low-to-moderate severity disturbance in regrown century-old forests altered trajectories of canopy rugosity. We conclude that structural complexity indicators display a more nuanced temporal and directional response to disturbance than conventional leaf area and cover indexes. Predicting what disturbance conditions modify trajectories of structural complexity remains critical to disturbance characterization and the inference of ecosystem functioning. [ABSTRACT FROM AUTHOR]

Carbon (C) cycling processes are particularly dynamic following disturbance, with initial responses often indicative of longer-term change. In northern Michigan, USA, we initiated the Forest Resilience Threshold Experiment (FoRTE) to identify the processes that sustain or lead to the decline of C cycling rates across multiple levels (0, 45, 65 and 85% targeted gross leaf area index loss) of disturbance severity and, in response, to separate disturbance types preferentially targeting large or small diameter trees. Simulating the effects of boring insects, we stem girdled > 3600 trees below diameter at breast height (DBH), immediately and permanently disrupting the phloem. Weekly DBH measurements of girdled and otherwise healthy trees (n > 700) revealed small but significant increases in daily aboveground wood net primary production (ANPPw) in the 65 and 85% disturbance severity treatments that emerged six weeks after girdling. However, we observed minimal change in end-of-season leaf area index and no significant differences in annual ANPPw among disturbance severities or between disturbance types, suggesting continued C fixation by girdled trees sustained stand-scale wood production in the first growing season after disturbance. We hypothesized higher disturbance severities would favor the growth of early successional species but observed no significant difference between early and middle to late successional species' contributions to ANPPw across the disturbance severity gradient. We conclude that ANPPw stability immediately following phloem disruption is dependent on the continued, but inevitably temporary, growth of phloem-disrupted trees. Our findings provide insight into the tree-to-ecosystem mechanisms supporting stand-scale wood production stability in the first growing season following a phloem-disrupting disturbance. [ABSTRACT FROM AUTHOR]

Structure–function relationships are central to many ecological paradigms. Chief among these is the linkage of net primary production (NPP) with species diversity and canopy structure. Using the National Ecological Observatory Network (NEON) as a subcontinental‐scale research platform, we examined how temperate‐forest NPP relates to several measures of site‐level canopy structure and tree species diversity. Novel multidimensional canopy traits describing structural complexity, most notably canopy rugosity, were more strongly related to site NPP than were species diversity measures and other commonly characterized canopy structural features. The amount of variation in site‐level NPP explained by canopy rugosity alone was 83%, which was substantially greater than that explained individually by vegetation area index (31%) or Shannon's index of species diversity (30%). Forests that were more structurally complex, had higher vegetation‐area indices, or were more diverse absorbed more light and used light more efficiently to power biomass production, but these relationships were most strongly tied to structural complexity. Implications for ecosystem modeling and management are wide ranging, suggesting structural complexity traits are broad, mechanistically robust indicators of NPP that, in application, could improve the prediction and management of temperate forest carbon sequestration. [ABSTRACT FROM AUTHOR]