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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.

Adaptations to different environments between closely related species can be important drivers of reproductive isolation during speciation due to either habitat isolation between species or reduced fitness in hybrids that possess suboptimal adaptations. Hybrid zones are useful natural arenas to explore how ecologically divergent species compete for habitat in sympatry and how possible differences in their habitat use may contribute to the speciation process. We investigated habitat selection by sister species of tree squirrels, the Douglas squirrel and the red squirrel, that have evolved in different forest types in allopatry and hybridize in a transitional forest. We first used genome-wide SNP data and admixture analyses to classify individuals into parental or hybrid classes. Next, we estimated home ranges with radio telemetry data and then used a novel ground-based lidar system to measure forest canopy structure of squirrel home ranges, midden sites, and marginally used forest habitat. We found hybrids consisting of multiple hybrid classes were intermixed with both parental species in the same forest with varying canopy structure complexity. On average, Douglas squirrels utilized forests with slightly greater structural complexity than either red squirrels or hybrids, while marginally used forests were the least structurally complex. Interestingly, hybrid squirrels were not relegated to marginal habitat and were successful in mating among each other and with both parental species. As such, our study suggests that prezygotic-ecological isolation and postzygotic-hybrid infertility, and postzygotic ecological inviability of hybrids are not strong barriers in the speciation process between Douglas squirrels and red squirrels. Closely related species that become geographically divided often encounter different environments and thus evolve different adaptations. Sometimes, these species meet again and produce hybrids. Hybrids often have lower fitness due to their inferior adaptations. We studied a pair of closely related squirrels that evolved in different environments and meet and produce hybrids in a secondary contact zone. Despite these species evolving in allopatry in very different forest types, they do not select different forest characteristics while in sympatry in the hybrid zone. Furthermore, hybrids do not show major differences in types of habitats that they choose and are capable of defending territories and reproducing among each other and with both parental species. In summary, prezygotic and postzygotic isolating mechanisms associated with habitat selection do not appear to have an important role in promoting species divergence.

The structural dynamics of forest canopies involve complex interactions among the abiotic environment, stand structure, species composition and disturbance regimes. How the re-arrangement of tree canopies in space and time affects forest aboveground net primary productivity (ANPP) remains poorly understood, however. In this study, we analyzed a long-term dataset from a temperate deciduous forest in Northern Michigan, USA, to investigate two primary objectives: 1) what abiotic and biotic factors influence canopy complexity and its inter-annual variability, and 2) the direct and indirect effects that abiotic, biotic and canopy complexity variables have on ANPP. We hypothesized that inter-annual variability in canopy complexity would be lower in high complexity canopies and that temporal variability in complexity metrics would be inversely related to ANPP. We found that canopy complexity was highest in more taxonomically diverse stands with high variability in tree diameters and in stands dominated by Populus tremuloides and Populus grandidentata. Canopy complexity was lowest in stands dominated by Quercus rubra and Pinus strobus, which also had lower ANPP. Stands with a high stem density had lower inter-annual variation in canopy complexity, exhibited more height growth and an increase in canopy open space, which in turn enhanced ANPP. Our results provide novel empirical evidence linking temporal stability in canopy complexity to ANPP, and suggest that variability in canopy complexity over time, in addition to the overall mean canopy complexity, may be important when considering drivers of forest carbon uptake.

Summary 1.Forest ecosystems are critical for the global regulation of carbon (C), a substantial portion of which is stored in aboveground biomass (AGB). While it is well understood that taxonomic and functional composition, stand structure, and environmental gradients influence spatial variation in AGB, the relative strengths of these drivers at landscape-scales has not been investigated in temperate forests. Furthermore, when biodiversity enhances C storage, it is unclear whether it is through mass-ratio effects (i.e., the dominant trait in communities regulates AGB) or through niche complementarity (i.e., increased AGB due to interspecific resource partitioning). 2.To address these mechanisms, we analyzed data from a census of 28,262 adult trees sampled across 900 ha of temperate deciduous forest in southwestern Pennsylvania. We used data on four key plant functional traits to determine if (1) there is a positive relationship between species diversity and AGB and (2) whether this is due to mass-ratio effects or niche complementarity. We also sought to (3) identify the physical stand structural attributes and topographic variables that influence AGB across this landscape. 3.We found AGB was positively related to species richness and negatively related to species evenness, albeit weakly, while functional diversity indices had neutral effects. AGB was enhanced in communities dominated by traits related to greater maximum tree height, deeper minimum rooting depths and larger seeds. Most importantly, areas with high AGB were dominated by Acer saccharum and Liriodendron tulipifera. Overall, these results support mass-ratio effects, with little evidence for niche complementarity. 4.Synthesis: Stand structure, topography, and species and functional composition, but not taxonomic or functional diversity, were found to be key drivers of AGB at landscape-scales (

Canopy structure influences forest productivity through its effects on the distribution of radiation and the light-induced changes in leaf physiological traits. Due to the difficulty of accessing and measuring forest canopies, few field-based studies have quantitatively linked these divergent scales of canopy functioning. The objective of our study was to investigate how canopy structure affects light profiles within a forest canopy and whether leaves of mature trees adjust morphologically and biochemically to the light environments characteristic of canopies with different structural complexity. We used a combination of light detection and ranging (LiDAR) data and hemispherical photographs to quantify canopy structure and light environments, respectively, and a telescoping pole to sample leaves. Leaf mass per area (LMA), nitrogen on an area basis (Narea) and chlorophyll on a mass basis (Chlmass) were measured in red maple (Acer rubrum), american beech (Fagus grandifolia), white pine (Pinus strobus), and northern red oak (Quercus rubra) at different heights in plots with similar leaf area index but contrasting canopy complexity (rugosity). We found that more complex canopies had greater porosity and reduced light variability in the midcanopy while total light interception was unchanged relative to less complex canopies. Leaf phenotypes of F. grandifolia, Q. rubra and P. strobus were more sun-acclimated in the midstory of structurally complex canopies while leaf phenotypes of A. rubrum were more shade-acclimated (lower LMA) in the upper canopy of more complex stands, despite no differences in total light interception. Broadleaf species showed further differences in acclimation with increased Narea and reduced Chlmass in leaves with higher LMA, while P. strobus showed no change in Narea and Chlmass with higher LMA. Our results provide new insight on how light distribution and leaf acclimation in mature trees might be altered when natural and anthropogenic disturbances cause structural changes in the canopy.

Moderate severity disturbances, which only kill a subset of canopy trees (e.g., via insects, pathogens, and windthrow), are increasingly widespread in North America, and can alter forest structure and production. Whether the net primary production (NPP) of forest stands differing in pre-disturbance site quality and composition respond similarly to moderate severity disturbance, however, is unknown, but critical to understanding the disturbance response dynamics of patchy landscapes. We experimentally disturbed three, 2-ha stands varying in pre-disturbance primary production and community composition, temporarily reducing live stand basal area by 38% to 66% through the stem girdling of all mature early successional aspen (Populus tremuloides Michx. and Populus grandidentata Michx.) and birch (Betula papyrifera Marshall). Disturbance significantly altered stand-scale physical and biological structure and prompted a similar decade-long pattern of wood NPP decline and recovery. All stands exhibited an initial reduction in wood NPP, followed by a recovery period and eventual return to pre-disturbance levels within eight years, with the most productive stand exhibiting an increase in primary production following recovery. Following wood NPP recovery, more biologically diverse forest canopies with higher leaf area indexes intercepted more light, and, consequently, had higher rates of wood NPP. We conclude that, despite substantial pre-disturbance differences in productivity and community composition, relative wood NPP recovery patterns can be similar, though long-term post-recovery primary production may trend higher in more productive and compositionally diverse stands. We suggest that improved mechanistic understanding of different forest ecosystems’ responses to disturbances remains critical to informing management decisions across diverse landscape mosaics.

The regrowing forests of eastern North America have been an important global C sink over the past 100+ years, but many are now transitioning into late succession. The consequences of this transition are unclear due to uncertainty around the C dynamics of old- growth forests. Canopy structural complexity (CSC) has been shown to be an important source of variability in C dynamics in younger forests (e.g., in productivity and resilience to disturbance), but its role in late-successional forests has not been widely addressed. We investigated patterns of CSC in two old-growth forest landscapes in the Upper Peninsula of Michigan, USA, to assess factors associated with CSC and its influence on productivity and disturbance resilience (to moderate-severity windstorm). CSC was quantified using a portable below-canopy LiDAR (PCL) system in 65 plots that also had long-term (50-70+ years). inventory data, which were used to quantify aboveground net primary productivity (ANPP), disturbance history, and stand characteristics. We found high and variable CSC relative to younger forests across a suite of PCL-derived metrics. Variation in CSC was driven by species composition and size structure, rather than disturbance history or site characteristics. Recent moderate severity wind disturbance decreased plot-scale CSC, but increased stand-scale variation in CSC. The strong positive correlation between CSC and productivity illustrated in younger forests was not present in undisturbed portions of these late-successional ecosystems. Moderate severity disturbance appeared to reestablish the positive link between CSC and productivity, but this relationship was scale and severity dependent. A positive CSC-productivity relationship was evident at the plot scale with low-severity, dispersed disturbance, but only at a patch scale in more severely disturbed areas. CSC does not appear to strongly correlate With variation in productivity in undisturbed old-growth forests, but may play a very important (and scale/severity-dependent) role in their response to disturbance. Understanding potential, drivers and consequences of CSC in late-successional forests will inform management focused on promoting complexity and old-growth conditions, and illustrate potential inipacts of such treatments on regional C dynamics.