North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

Visions of Restoration in Fire-Adapted Forest Landscapes: Lessons from the Collaborative Forest Landscape Restoration Program.

Collaborative approaches to natural resource management are becoming increasingly common on public lands. Negotiating a shared vision for desired conditions is a fundamental task of collaboration and serves as a foundation for developing management objectives and monitoring strategies. We explore the complex socio-ecological processes involved in developing a shared vision for collaborative restoration of fire-adapted forest landscapes. To understand participant perspectives and experiences, we analyzed interviews with 86 respondents from six collaboratives in the western U.S., part of the Collaborative Forest Landscape Restoration Program established to encourage collaborative, science-based restoration on U.S. Forest Service lands. Although forest landscapes and group characteristics vary considerably, collaboratives faced common challenges to developing a shared vision for desired conditions. Three broad categories of challenges emerged: meeting multiple objectives, collaborative capacity and trust, and integrating ecological science and social values in decision-making. Collaborative groups also used common strategies to address these challenges, including some that addressed multiple challenges. These included use of issue-based recommendations, field visits, and landscape-level analysis; obtaining support from local agency leadership, engaging facilitators, and working in smaller groups (sub-groups); and science engagement. Increased understanding of the challenges to, and strategies for, developing a shared vision of desired conditions is critical if other collaboratives are to learn from these efforts.

A cross-continental comparison of plant and beetle responses to retention of forest patches during timber harvest.

Timber harvest can adversely affect forest biota. Recent research and application suggest that retention of mature forest elements (retention forestry), including unharvested patches (or aggregates) within larger harvested units, can benefit biodiversity compared to clearcutting. However, it is unclear whether these benefits can be generalized among the diverse taxa and biomes in which retention forestry is practiced. Lack of comparability in methods for sampling and analyzing responses to timber harvest and edge creation presents a challenge to synthesis. We used a consistent methodology (similarly spaced plots or traps along transects) to investigate responses of vascular plants and ground-active beetles to aggregated retention at replicate sites in each of four temperate and boreal forest types on three continents: Douglas-fir forests in Washington, USA; aspen forests in Minnesota, USA; spruce forests in Sweden; and wet eucalypt forests in Tasmania, Australia. We assessed (1) differences in local (plot-scale) species richness and composition between mature (intact) and regenerating (previously harvested) forest; (2) the lifeboating function of aggregates (capacity to retain species of unharvested forest); and whether intact forests and aggregates (3) are susceptible to edge effects and (4) influence the adjacent regenerating forest. Intact and harvested forests differed in composition but not richness of plants and beetles. The magnitude of this difference was generally similar among regions, but there was considerable heterogeneity of composition within and among replicate sites. Aggregates within harvest units were effective at lifeboating for both plant and beetle communities. Edge effects were uncommon even within the aggregates. In contrast, effects of forest influence on adjacent harvested areas were common and as strong for aggregates as for larger blocks of intact forest. Our results provide strong support for the widespread application of aggregated retention in boreal and temperate forests. The consistency of pattern in four very different regions of the world suggests that, for forest plants and beetles, responses to aggregated retention are likely to apply more widely. Our results suggest that through strategic placement of aggregates, it is possible to maintain the natural heterogeneity and biodiversity of mature forests managed for multiple objectives.

Spatial aspects of tree mortality strongly differ between young and old-growth forests.

Rates and spatial patterns of tree mortality are predicted to change during forest structural development. In young forests, mortality should be primarily density dependent due to competition for light, leading to an increasingly spatially uniform pattern of surviving trees. In contrast, mortality in old-growth forests should be primarily caused by contagious and spatially autocorrelated agents (e.g., insects, wind), causing spatial aggregation of surviving trees to increase through time. We tested these predictions by contrasting a three-decade record of tree mortality from replicated mapped permanent plots located in young (< 60-year-old) and old-growth (> 300-year-old) Abies amabilis forests. Trees in young forests died at a rate of 4.42% per year, whereas trees in old-growth forests died at 0.60% per year. Tree mortality in young forests was significantly aggregated, strongly density dependent, and caused live tree patterns to become more uniform through time. Mortality in old-growth forests was spatially aggregated, but was density independent and did not change the spatial pattern of surviving trees. These results extend current theory by demonstrating that density-dependent competitive mortality leading to increasingly uniform tree spacing in young forests ultimately transitions late in succession to a more diverse tree mortality regime that maintains spatial heterogeneity through time.

Spatially nonrandom tree mortality and ingrowth maintain equilibrium pattern in an old-growth Pseudotsuga-Tsuga forest.

Mortality processes in old-growth forests are generally assumed to be driven by gap-scale disturbance, with only a limited role ascribed to density-dependent mortality, but these assumptions are rarely tested with data sets incorporating repeated measurements. Using a 12-ha spatially explicit plot censused 13 years apart in an approximately 500-year-old Pseudotsuga-Tsuga forest, we demonstrate significant density-dependent mortality and spatially aggregated tree recruitment. However, the combined effect of these strongly nonrandom demographic processes was to maintain tree patterns in a state of dynamic equilibrium. Density-dependent mortality was most pronounced for the dominant late-successional species, Tsuga heterophylla. The long-lived, early-seral Pseudotsuga menziesii experienced an annual stem mortality rate of 0.84% and no new recruitment. Late-seral species Tsuga and Abies amabilis had nearly balanced demographic rates of ingrowth and mortality. The 2.34% mortality rate for Taxus brevifolia was higher than expected, notably less than ingrowth, and strongly affected by proximity to Tsuga. Large-diameter Tsuga structured both the regenerating conspecific and heterospecific cohorts with recruitment of Tsuga and Abies unlikely in neighborhoods crowded with large-diameter competitors (P < 0.001). Density-dependent competitive interactions strongly shape forest communities even five centuries after stand initiation, underscoring the dynamic nature of even equilibrial old-growth forests.

Masting is uncommon in trees that depend on mutualist dispersers in the context of global climate and fertility gradients.

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands.

Continent-wide tree fecundity driven by indirect climate effects.

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

Potential site productivity influences the rate of forest structural development.

Development and maintenance of structurally complex forests in landscapes formerly managed for timber production is an increasingly common management objective. It has been postulated that the rate of forest structural development increases with site productivity. We tested this hypothesis for Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forests using a network of permanent study plots established following complete timber harvest of the original old-growth forests. Forest structural development was assessed by comparing empirical measures of live tree structure to published values for Douglas-fir forests spanning a range of ages and structural conditions. The rate of forest structural development--resilience--exhibited a positive relationship with site index, a measure of potential site productivity. Density of shade-intolerant conifers declined in all study stands from an initial range of 336-4068 trees/ha to a range of 168-642 trees/ha at the most recent measurement. Angiosperm tree species declined from an initial range of 40-371 trees/ha to zero in seven of the nine plots in which they were present. Trends in shade-tolerant tree density were complex: density ranged from 0 to 575 trees/ha at the first measurement and was still highly variable (25-389 trees/ha) at the most recent measurement. Multivariate analysis identified the abundance of hardwood tree species as the strongest compositional trend apparent over the study period. However, structural variables showed a strong positive association with increasing shade-tolerant basal area and little or no association with abundance of hardwood species. Thus, while tree species succession and forest structural development occur contemporaneously, they are not equivalent processes, and their respective rates are not necessarily linearly related. The results of this study support the idea that silvicultural treatments to accelerate forest structural development should be concentrated on lower productivity sites when the management objective is reserve-wide coverage of structurally complex forests. Alternatively, high-productivity sites should be prioritized for restoration treatments when the management objective is to develop structurally complex forests on a portion of the landscape.

Earth Observation Networks (EONs): Finding the Right Balance.

Earth observation networks (EONs) are an emerging, surveillance-based approach to environmental monitoring and research that are fundamentally different than traditional question-driven, experimentally designed approaches. There is an urgent need to find an optimal balance between these approaches and to develop new integrated initiatives that take advantage of key features of them both.

Average Stand Age from Forest Inventory Plots Does Not Describe Historical Fire Regimes in Ponderosa Pine and Mixed-Conifer Forests of Western North America.

Quantifying historical fire regimes provides important information for managing contemporary forests. Historical fire frequency and severity can be estimated using several methods; each method has strengths and weaknesses and presents challenges for interpretation and verification. Recent efforts to quantify the timing of historical high-severity fire events in forests of western North America have assumed that the "stand age" variable from the US Forest Service Forest Inventory and Analysis (FIA) program reflects the timing of historical high-severity (i.e. stand-replacing) fire in ponderosa pine and mixed-conifer forests. To test this assumption, we re-analyze the dataset used in a previous analysis, and compare information from fire history records with information from co-located FIA plots. We demonstrate that 1) the FIA stand age variable does not reflect the large range of individual tree ages in the FIA plots: older trees comprised more than 10% of pre-stand age basal area in 58% of plots analyzed and more than 30% of pre-stand age basal area in 32% of plots, and 2) recruitment events are not necessarily related to high-severity fire occurrence. Because the FIA stand age variable is estimated from a sample of tree ages within the tree size class containing a plurality of canopy trees in the plot, it does not necessarily include the oldest trees, especially in uneven-aged stands. Thus, the FIA stand age variable does not indicate whether the trees in the predominant size class established in response to severe fire, or established during the absence of fire. FIA stand age was not designed to measure the time since a stand-replacing disturbance. Quantification of historical "mixed-severity" fire regimes must be explicit about the spatial scale of high-severity fire effects, which is not possible using FIA stand age data.

Preserving Biodiversity: Species, Ecosystems, or Landscapes?

Efforts to preserve biological diversity must focus increasingly at the ecosystem level because of the immense number of species, the majority of which are currently unknown. An ecosystem approach is also the only way to conserve processes and habitats (such as forest canopies, belowground habitats, and hyporheic zones) that, with their constituent species, are poorly known. Continued concern with species is essential, however. Landscape-level issues also need much greater attention. Designing an appropriate system of habitat reserves is one landscape-level concern. Understanding and appropriately manipulating the landscape matrix is at least equal in importance to reserves issues, however, since the matrix itself is important in maintaining diversity, influences the effectiveness of reserves, and controls landscape connectivity.

Vegetation Responses to Edge Environments in Old-Growth Douglas-Fir Forests.

Forest edges created by dispersed-patch clear-cutting have become a conspicuous landscape feature in western North America, but the effects of edge on forest structure and function are still poorly understood. In this paper we describe responses of stocking density, growth, mortality, and regeneration for three conifer species from the clear-cut edge into the interior of old-growth forest patches adjacent to 10-15 yr old clearcuts in southern Washington and central Oregon. The significance of edge effects for each variable was tested through a single-factor (distance) analysis of variance (F test). Relationships between these variables and depth-of-edge influence (i.e., edge width) on old-growth forest were characterized by nonlinear regression models. Near the edge (forest-clearcut boundary line), the old-growth forest has (1) reduced stocking density, as measured by canopy cover, number of stems per hectare, and basal area; (2) increased growth rates of dominant Douglas-fir (Pseudotsuga menziesii) and western hemlock (Tsuga heterophylla), as calculated by an index of relative growth rate; (3) elevated rates of tree mortality, as measured by standing dead and down trees (snags and logs); and (4) greater numbers of Douglas-fir and western hemlock seedlings (@<100 cm tall) and saplings (101-200 cm) but fewer of Pacific silver fir (Abies amabilis). The depth-of-edge influence, when calculated as the point along the clearcut-forest gradient at which a given variable has returned to a condition representing 2/3 of the interior forest environment, ranged from 16 to 137 m for variables related to distance from the edge. The amount of a square forest patch affected by edge decreased as patch size increased and varied greatly with the depth-of-edge influence. With increasing concerns about organisms and processes that require interior forest habitat, determining the area of residual forest influenced by adjacent clearcut is critical to current and future resource management. Responses of additional biological variables must be explored and information on edge phenomena should be extended to the scale of landscapes.

Aging in Pacific Northwest forests: a selection of recent research.

This paper provides an overview of the work reported at a symposium on age-related changes in the structure and function of forests in the United States Pacific Northwest. Some of the work presented at this meeting is reported in the peer-reviewed papers comprising this journal issue. Age-related changes in leaf structure, CO2 assimilation rate, stable carbon isotope ratio, nitrogen concentration and stomatal limitation were demonstrated at many organizational scales. At larger scales, age-related changes were reported in canopy structure and light profile, stand productivity, tree mortality and respiration. These data raise new questions about the potential interaction among the structural and functional changes in aging forests, and indicate many avenues for future research concerning tree growth and ecosystem functioning.

Can retention forestry help conserve biodiversity? A meta-analysis.

Industrial forestry typically leads to a simplified forest structure and altered species composition. Retention of trees at harvest was introduced about 25 years ago to mitigate negative impacts on biodiversity, mainly from clearcutting, and is now widely practiced in boreal and temperate regions. Despite numerous studies on response of flora and fauna to retention, no comprehensive review has summarized its effects on biodiversity in comparison to clearcuts as well as un-harvested forests. Using a systematic review protocol, we completed a meta-analysis of 78 studies including 944 comparisons of biodiversity between retention cuts and either clearcuts or un-harvested forests, with the main objective of assessing whether retention forestry helps, at least in the short term, to moderate the negative effects of clearcutting on flora and fauna. Retention cuts supported higher richness and a greater abundance of forest species than clearcuts as well as higher richness and abundance of open-habitat species than un-harvested forests. For all species taken together (i.e. forest species, open-habitat species, generalist species and unclassified species), richness was higher in retention cuts than in clearcuts. Retention cuts had negative impacts on some species compared to un-harvested forest, indicating that certain forest-interior species may not survive in retention cuts. Similarly, retention cuts were less suitable for some open-habitat species compared with clearcuts. Positive effects of retention cuts on richness of forest species increased with proportion of retained trees and time since harvest, but there were not enough data to analyse possible threshold effects, that is, levels at which effects on biodiversity diminish. Spatial arrangement of the trees (aggregated vs. dispersed) had no effect on either forest species or open-habitat species, although limited data may have hindered our capacity to identify responses. Results for different comparisons were largely consistent among taxonomic groups for forest and open-habitat species, respectively. Synthesis and applications. Our meta-analysis provides support for wider use of retention forestry since it moderates negative harvesting impacts on biodiversity. Hence, it is a promising approach for integrating biodiversity conservation and production forestry, although identifying optimal solutions between these two goals may need further attention. Nevertheless, retention forestry will not substitute for conservation actions targeting certain highly specialized species associated with forest-interior or open-habitat conditions. Our meta-analysis provides support for wider use of retention forestry since it moderates negative harvesting impacts on biodiversity. Hence, it is a promising approach for integrating biodiversity conservation and production forestry, although identifying optimal solutions between these two goals may need further attention. Nevertheless, retention forestry will not substitute for conservation actions targeting certain highly specialized species associated with forest-interior or open-habitat conditions.

Interacting factors driving a major loss of large trees with cavities in a forest ecosystem.

Large trees with cavities provide critical ecological functions in forests worldwide, including vital nesting and denning resources for many species. However, many ecosystems are experiencing increasingly rapid loss of large trees or a failure to recruit new large trees or both. We quantify this problem in a globally iconic ecosystem in southeastern Australia--forests dominated by the world's tallest angiosperms, Mountain Ash (Eucalyptus regnans). Tree, stand and landscape-level factors influencing the death and collapse of large living cavity trees and the decay and collapse of dead trees with cavities are documented using a suite of long-term datasets gathered between 1983 and 2011. The historical rate of tree mortality on unburned sites between 1997 and 2011 was >14% with a mortality spike in the driest period (2006-2009). Following a major wildfire in 2009, 79% of large living trees with cavities died and 57-100% of large dead trees were destroyed on burned sites. Repeated measurements between 1997 and 2011 revealed no recruitment of any new large trees with cavities on any of our unburned or burned sites. Transition probability matrices of large trees with cavities through increasingly decayed condition states projects a severe shortage of large trees with cavities by 2039 that will continue until at least 2067. This large cavity tree crisis in Mountain Ash forests is a product of: (1) the prolonged time required (>120 years) for initiation of cavities; and (2) repeated past wildfires and widespread logging operations. These latter factors have resulted in all landscapes being dominated by stands ≤72 years and just 1.16% of forest being unburned and unlogged. We discuss how the features that make Mountain Ash forests vulnerable to a decline in large tree abundance are shared with many forest types worldwide.