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Recent ecological research suggests that, in general, mixtures are more resistant to insect herbivores and pathogens than monocultures. However, we know little about mixtures with non-native trees, where enemy release could lead to patterns that differ from commonly observed relationships among native species. This becomes particularly relevant when considering that adaptation strategies to climate change increasingly promote a larger share of non-native tree species, such as North American Douglas fir in Central Europe. We studied leaf damage on European beech (Fagus sylvatica) saplings and mature trees across a wide range of site conditions in monocultures and mixtures with phylogenetically distant conifers native Norway spruce (Picea abies) and non-native Douglas fir (Pseudotsuga menziesii). We analyzed leaf herbivory and pathogen damage in relation to tree diversity and composition effects, as well as effects of environmental factors and plant characteristics. We observed lower sapling herbivory and tree sucking damage on beech in non-native Douglas fir mixtures than in beech monocultures, probably due to a lower herbivore diversity on Douglas fir trees, and higher pathogen damage on beech saplings in Norway spruce than Douglas fir mixtures, possibly because of higher canopy openness. Our findings suggest that for low diversity gradients, tree diversity effects on leaf damage can strongly depend on tree species composition, in addition to modifications caused by feeding guild and tree ontogeny. Moreover, we found that nutrient capacity modulated the effects of tree diversity, composition, and environmental factors, with different responses in sites with low or high nutrient capacity. The existence of contrasting diversity effects based on tree species composition provides important information on our understanding of the relationships between tree diversity and plant-herbivore interactions in light of non-native tree species introductions. Especially with recent Norway spruce die-off, the planting of Douglas fir as replacement is likely to strongly increase in Central Europe. Our findings suggest that mixtures with Douglas fir could benefit the survival or growth rates of beech saplings and mature trees due to lower leaf damage, emphasizing the need to clearly identify and compare the potential benefits and ecological trade-offs of non-native tree species in forest management under ongoing environmental change., (© 2022 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Tree masting is one of the most intensively studied ecological processes. It affects nutrient fluxes of trees, regeneration dynamics in forests, animal population densities, and ultimately influences ecosystem services. Despite a large volume of research focused on masting, its evolutionary ecology, spatial and temporal variability, and environmental drivers are still matter of debate. Understanding the proximate and ultimate causes of masting at broad spatial and temporal scales will enable us to predict tree reproductive strategies and their response to changing environment. Here we provide broad spatial (distribution range-wide) and temporal (century) masting data for the two main masting tree species in Europe, European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) H. Karst.). We collected masting data from a total of 359 sources through an extensive literature review and from unpublished surveys. The data set has a total of 1,747 series and 18,348 yearly observations from 28 countries and covering a time span of years 1677-2016 and 1791-2016 for beech and spruce, respectively. For each record, the following information is available: identification code; species; year of observation; proxy of masting (flower, pollen, fruit, seed, dendrochronological reconstructions); statistical data type (ordinal, continuous); data value; unit of measurement (only in case of continuous data); geographical location (country, Nomenclature of Units for Territorial Statistics NUTS-1 level, municipality, coordinates); first and last record year and related length; type of data source (field survey, peer reviewed scientific literature, gray literature, personal observation); source identification code; date when data were added to the database; comments. To provide a ready-to-use masting index we harmonized ordinal data into five classes. Furthermore, we computed an additional field where continuous series with length >4 yr where converted into a five classes ordinal index. To our knowledge, this is the most comprehensive published database on species-specific masting behavior. It is useful to study spatial and temporal patterns of masting and its proximate and ultimate causes, to refine studies based on tree-ring chronologies, to understand dynamics of animal species and pests vectored by these animals affecting human health, and it may serve as calibration-validation data for dynamic forest models., (© 2017 by the Ecological Society of America.)

Detailed understanding of forest disturbance interactions is needed for effective forecasting, modelling, and management. Insect outbreaks are a significant forest disturbance that alters forest structure as well as the distribution and connectivity of combustible fuels at broad spatial scales. The effect of insect outbreaks on fire activity is an important but contentious issue with significant policy consequences. The eastern spruce budworm (Choristoneura fumiferana) is a native defoliating insect in eastern North America whose periodic outbreaks create large patches of dead fir and spruce trees. Of particular concern to fire and forest managers is whether these patches represent an increased fire risk, if so, for how long, and how the relationship between defoliation and fire risk varies through space and time. Previous work suggests a temporary increase in flammability in budworm-killed forests, but regional and seasonal variability in these relationships has not been examined. Using an extensive database on historical lightning-caused fire ignitions and spruce budworm defoliation between 1963 and 2000, we assess the relative importance of cumulative defoliation and fire weather on the probability of ignition in Ontario, Canada. We modeled fire ignition using a generalized additive logistic regression model that accounts for temporal autocorrelation in fire weather. We compared two ecoregions in eastern Ontario (Abitibi Plains) and western Ontario (Lake of the Woods) that differ in terms of climate, geomorphology, and forest composition. We found that defoliation has the potential to both increase and decrease the probability of ignition depending on the time scale, ecoregion, and season examined. Most importantly, we found that lagged spruce budworm defoliation (8-10 yr) increases the risk of fire ignition whereas recent defoliation (1 yr) can decrease this risk. We also found that historical defoliation has a greater influence on ignition risk during the spring than during the summer fire season. Given predicted increases in forest insect activity due to global change, these results represent important information for fire management agencies that can be used to refine existing models of fire risk., (© 2016 by the Ecological Society of America.)

The severity and spatial extent of bark-beetle outbreaks substantially increased in recent decades worldwide. The ongoing controversy about natural forest recovery after these outbreaks highlights the need for individual-based long-term studies, which disentangle processes driving forest regeneration. However, such studies have been lacking. To fill this gap, we followed the fates of 2,552 individual seedlings for 12 years after a large-scale bark-beetle outbreak that caused complete canopy dieback in mountain Norway spruce (Picea abies) forests in southeast Germany. We explore the contribution of advance, disturbance-related, and post-disturbance regeneration to forest recovery. Most seedlings originated directly within the three-year dieback of canopy trees induced by bark-beetle outbreak. After complete canopy dieback, the establishment of new seedlings was minimal. Surprisingly, advance regeneration formed only a minor part of all regeneration. However, because it had the highest survival rate, its importance increased over time. The most important factor influencing the survival of seedlings after disturbance was their height. Survival was further modified by microsite: seedlings established on dead wood survived best, whereas almost all seedlings surrounded by graminoids died. For 5 cm tall seedlings, annual mortality ranged from 20 to 50% according to the rooting microsite. However, for seedlings taller than 50 cm, annual mortality was below 5% at all microsites. While microsite modified seedling mortality, it did not affect seedling height growth. A model of regeneration dynamics based on short-term observations accurately predicts regeneration height growth, but substantially underestimates mortality rate, thus predicting more surviving seedlings than were observed. We found that P. abies forests were able to regenerate naturally even after severe bark-beetle outbreaks owing to advance and particularly disturbance-related regeneration. This, together with microsite-specific mortality, yields structurally and spatially diverse forests. Our study thus highlights the so far unrecognized importance of disturbance-related regeneration for stand recovery after bark-beetle outbreaks., (© 2016 by the Ecological Society of America.)

Increasing temperatures have resulted in reduced growth and increased tree mortality across large areas of western North American forests. We use tree-ring isotope chronologies (δ 13 C and δ 18 O) from live and dead trees from four locations in south-central Alaska, USA, to test whether white spruce trees killed by recent spruce beetle (Dendroctonus rufipennis Kirby) outbreaks showed evidence of drought stress prior to death. Trees that were killed were more sensitive to spring/summer temperature and/or precipitation than trees that survived. At two of our sites, we found greater correlations between the δ 13 C and δ 18 O chronologies and spring/summer temperatures in dead trees than in live trees, suggesting that trees that are more sensitive to temperature-induced drought stress are more likely to be killed. At one site, the difference between δ 13 C in live and dead trees was related to winter/spring precipitation, with dead trees showing stronger correlations between δ 13 C and precipitation, again suggesting increased water stress in dead trees. At all sites where δ 18 O was measured, δ 18 O chronologies showed the greatest difference in climate response between live and dead groups, with δ 18 O in live trees correlating more strongly with late winter precipitation than dead trees. Our results indicate that sites where trees are already sensitive to warm or dry early growing-season conditions experienced the most beetle-kill, which has important implications for forecasting future mortality events in Alaska., (© 2016 by the Ecological Society of America.)

Recent large and severe outbreaks of native bark beetles have raised concern among the general public and land managers about potential for amplified fire activity in western North America. To date, the majority of studies examining bark beetle outbreaks and subsequent fire severity in the U.S. Rocky Mountains have focused on outbreaks of mountain pine beetle (MPB; Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests, but few studies, particularly field studies, have addressed the effects of the severity of spruce beetle (Dendroctonus rufipennis Kirby) infestation on subsequent fire severity in subalpine Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests. In Colorado, the annual area infested by spruce beetle outbreaks is rapidly rising, while MPB outbreaks are subsiding; therefore understanding this relationship is of growing importance. We collected extensive field data in subalpine forests in the eastern San Juan Mountains, southwestern Colorado, USA, to investigate whether a gray-stage (< 5 yr from outbreak to time of fire) spruce beetle infestation affected fire severity. Contrary to the expectation that bark beetle infestation alters subsequent fire severity, correlation and multivariate generalized linear regression analysis revealed no influence of pre-fire spruce beetle severity on nearly all field or remotely sensed measurements of fire severity. Findings were consistent across moderate and extreme burning conditions. In comparison to severity of the pre-fire beetle outbreak, we found that topography, pre-outbreak basal area, and weather conditions exerted a stronger effect on fire severity. Our finding that beetle infestation did not alter fire severity is consistent with previous retrospective studies examining fire activity following other bark beetle outbreaks and reiterates the overriding influence of climate that creates conditions conducive to large, high-severity fires in the subalpine zone of Colorado. Both bark beetle outbreaks and wildfires have increased autonomously due to recent climate variability, but this study does not support the expectation that post-beetle outbreak forests will alter fire severity, a result that has important implications for management and policy decisions.

Increment cores from the boreal forest have long been used to reconstruct past climates. However, in recent years, numerous studies have revealed a deterioration of the correlation between temperature and tree growth that is commonly referred to as divergence. In the Brooks Range of northern Alaska, USA, studies of white spruce (Picea glauca) revealed that trees in the west generally showed positive growth trends, while trees in the central and eastern Brooks Range showed mixed and negative trends during late 20th century warming. The growing season climate of the eastern Brooks Range is thought to be drier than the west. On this basis, divergent tree growth in the eastern Brooks Range has been attributed to drought stress. To investigate the hypothesis that drought-induced stomatal closure can explain divergence in the Brooks Range, we synthesized all of the Brooks Range white spruce data available in the International Tree Ring Data Bank (ITRDB) and collected increment cores from our primary sites in each of four watersheds along a west-to-east gradient near the Arctic treeline. For cores from our sites, we measured ring widths and calculated carbon isotope discrimination (δ13C), intrinsic water-use efficiency (iWUE), and needle intercellular CO2 concentration (C(i)) from δ13C in tree-ring alpha-cellulose. We hypothesized that trees exhibiting divergence would show a corresponding decline in δ13C, a decline in C(i), and a strong increase in iWUE. Consistent with the ITRDB data, trees at our western and central sites generally showed an increase in the strength of the temperature-growth correlation during late 20th century warming, while trees at our eastern site showed strong divergence. Divergent tree growth was not, however, associated with declining δ13C. Meanwhile, estimates of C(i) showed a strong increase at all of our study sites, indicating that more substrate was available for photosynthesis in the early 21st than in the early 20th century. Our results, which are corroborated by measurements of xylem sap flux density, needle gas exchange, and measurements of growth and δ13C along moisture gradients within each watershed, suggest that drought-induced stomatal closure is probably not the cause of 20th century divergence in the Brooks Range.

Mast seeding involves the episodic and synchronous production of large seed crops by perennial plants. The predator satiation hypothesis proposes that mast seeding maximizes seed escape because seed predators consume a decreasing proportion of available seeds with increasing seed production. However, the seed escape benefits of masting depend not only on whether predators are satiated at high levels of seed production, but also on the shape of their functional response (type II vs. type III), and the actual proportion of available seeds that they consume at different levels of seed production. North American red squirrels (Tamiasciurus hudsonicus) are the primary vertebrate predator of white spruce (Picea glauca) mast seed crops in many boreal regions because they hoard unopened cones in underground locations, preempting the normal sequence of cone opening, seed dispersal, and seed germination. We document the functional response of cone-hoarding by red squirrels across three non-mast years and one mast year by estimating the number of cones present in the territories of individual red squirrels and the proportion of these cones that they hoarded each autumn. Even though red squirrels are not constrained by the ingestive and on-body (fat reserves) energy reserve limitations experienced by animals that consume seeds directly, most squirrels hoarded < 10% of the cones present on their territories under mast conditions. Cone availability during non-mast years also reached levels that satiated the hoarding activity of red squirrels; however, this occurred only on the highest-quality territories. Squirrels switched to mushroom-hoarding when cone production was low and mushrooms were abundant. This resulted in type III functional response whereby the proportional harvest of cones was highest at levels of cone availability that were intermediate within non-mast years. Overall, more cones escaped squirrel cone-hoarding during a mast event than when cone production was low in non-mast years, which supports the predator satiation hypothesis. However, the highly variable seed escape in non-mast years may help to explain why all spruce cone production is not concentrated into fewer, larger, mast years.

Black spruce (Picea mariana) is the most abundant tree species in the boreal biome, but little is known about how climate warming may change recruitment in peatlands, especially those affected by permafrost thaw. We used results from a seven-year study in northern Manitoba, Canada, to address the following questions: (1) What is the relative importance of early life history transitions on P. mariana recruitment? (2) How are these transitions mediated by biological and environmental factors, including competition, facilitation, disease, herbivory, water table depth, and soil nutrients? (3) Do interactions among these factors create additional recruitment limitations beyond those imposed by environmental factors changing with climate warming, such as hydrology? Seed rain was measured over six years on forested permafrost plateaus and in neighboring collapse scar bogs. Seed germination and seedling survival and growth were measured over 4-5 years in collapse scars and assessed across a three-level water table treatment. Survival and growth experiments examined additional combinations of above- and belowground vascular plant competition and fertilizer addition. Results showed that failure of germination and survival on growing moss surfaces and reduced survival of seedlings in wetter microsites were primary constraints. Seed influx was significantly lower in collapse scars but likely did not limit recruitment. Biological and environmental factors mediating these life history transitions also differed in relative importance, and interactions among them tended to amplify recruitment limitation. Seedling survival was most strongly controlled by fast-growing mosses in wet microsites but also was influenced by apparent drowning in wet plots, herbivory, and loss of foliage caused by a fungal pathogen. Seedling growth was strongly controlled by water table depth, nutrient and competition levels, and fungal pathogens. Multiple, interacting factors will affect P. mariana establishment in boreal peatlands during climate warming. Generalizations about recruitment relying on few environmental gradients sensitive to climate change, such as water table, may therefore not fully capture the complexities of establishment.

Two forest management objectives being debated in the context of federally managed landscapes in the U.S. Pacific Northwest involve a perceived trade-off between fire restoration and carbon sequestration. The former strategy would reduce fuel (and therefore C) that has accumulated through a century of fire suppression and exclusion which has led to extreme fire risk in some areas. The latter strategy would manage forests for enhanced C sequestration as a method of reducing atmospheric CO2 and associated threats from global climate change. We explored the trade-off between these two strategies by employing a forest ecosystem simulation model, STANDCARB, to examine the effects of fuel reduction on fire severity and the resulting long-term C dynamics among three Pacific Northwest ecosystems: the east Cascades ponderosa pine forests, the west Cascades western hemlock-Douglas-fir forests, and the Coast Range western hemlock-Sitka spruce forests. Our simulations indicate that fuel reduction treatments in these ecosystems consistently reduced fire severity. However, reducing the fraction by which C is lost in a wildfire requires the removal of a much greater amount of C, since most of the C stored in forest biomass (stem wood, branches, coarse woody debris) remains unconsumed even by high-severity wildfires. For this reason, all of the fuel reduction treatments simulated for the west Cascades and Coast Range ecosystems as well as most of the treatments simulated for the east Cascades resulted in a reduced mean stand C storage. One suggested method of compensating for such losses in C storage is to utilize C harvested in fuel reduction treatments as biofuels. Our analysis indicates that this will not be an effective strategy in the west Cascades and Coast Range over the next 100 years. We suggest that forest management plans aimed solely at ameliorating increases in atmospheric CO2 should forgo fuel reduction treatments in these ecosystems, with the possible exception of some east Cascades ponderosa pine stands with uncharacteristic levels of understory fuel accumulation. Balancing a demand for maximal landscape C storage with the demand for reduced wildfire severity will likely require treatments to be applied strategically throughout the landscape rather than indiscriminately treating all stands.

We measured soil climate and the turbulent fluxes of CO2, H2O, heat, and momentum on short towers (2 m) in a 160-yr-old boreal black spruce forest in Manitoba, Canada. Two distinct land cover types were studied: a Sphagnum-dominated wetland, and a feathermoss (Pleurozium and Hylocomium)-dominated upland, both lying within the footprint of a 30-m tower, which has measured whole-forest carbon exchange since 1994. Peak summertime uptake of CO2, was higher in the wetland than for the forest as a whole due to the influence of deciduous shrubs. Soil respiration rates in the wetland were approximately three times larger than in upland soils, and 30% greater than the mean of the whole forest, reflecting decomposition of soil organic matter. Soil respiration rates in the wetland were regulated by soil temperature, which was in turn influenced by water table depth through effects on soil heat capacity and conductivity. Warmer soil temperatures and deeper water tables favored increased heterotrophic respiration. Wetland drainage was limited by frost during the first half of the growing season, leading to high, perched water tables, cool soil temperatures, and much lower respiration rates than observed later in the growing season. Whole-forest evapotranspiration increased as water tables dropped, suggesting that photosynthesis in this forest was rarely subject to water stress. Our data indicate positive feedback between soil temperature, seasonal thawing, heterotrophic respiration, and evapotranspiration. As a result, climate warming could cause covariant changes in soil temperature and water table depths that may stimulate photosynthesis and strongly promote efflux of CO2 from peat soils in boreal wetlands.

Long-term forest productivity decline in boreal forests has been extensively studied in the last decades, yet its causes are still unclear. Soil conditions associated with soil organic matter accumulation are thought to be responsible for site productivity decline. The objectives of this study were to determine if paludification of boreal soils resulted in reduced forest productivity, and to identify changes in the physical and chemical properties of soils associated with reduction in productivity. We used a chronosequence of 23 black spruce stands ranging in postfire age from 50 to 2350 years and calculated three different stand productivity indices, including site index. We assessed changes in forest productivity with time using two complementary approaches: (1) by comparing productivity among the chronosequence stands and (2) by comparing the productivity of successive cohorts of trees within the same stands to determine the influence of time independently of other site factors. Charcoal stratigraphy indicates that the forest stands differ in their fire history and originated either from high- or low-severity soil burns. Both chronosequence and cohort approaches demonstrate declines in black spruce productivity of 50-80% with increased paludification, particularly during the first centuries after fire. Paludification alters bryophyte abundance and succession, increases soil moisture, reduces soil temperature and nutrient availability, and alters the vertical distribution of roots. Low-severity soil burns significantly accelerate rates of paludification and productivity decline compared with high-severity fires and ultimately reduce nutrient content in black spruce needles. The two combined approaches indicate that paludification can be driven by forest succession only, independently of site factors such as position on slope. This successional paludification contrasts with edaphic paludification, where topography and drainage primarily control the extent and rate of paludification. At the landscape scale, the fire regime (frequency and severity) controls paludification and forest productivity through its effect on soil organic layers. Implications for global carbon budgets and sustainable forestry are discussed.

Coarse woody debris (CWD) is a key habitat for many species in forest ecosystems. To ensure the long-term survival of such species, forest management regimes must include measures that promote dead wood dynamics similar to those of natural forests. Thus, information on CWD dynamics under natural conditions is required, including data pertaining to the underlying agents of disturbance. This study examines modes of mortality, decay rates, and temporal patterns in the availability of Picea abies logs in a Swedish old-growth forest affected by internal, small-scale disturbance. All 684 logs in a 6.6-ha plot were mapped and classified into one of six decay classes. Logs in the early stages of decay were examined for the presence of heart-rot fungi. Six years later all logs were re-inventoried, including newly formed logs. Matrix models based on the transition rates between decay classes showed that it took about 60 years for 90% of the logs to decay beyond class 6 (a deformed trunk with soft wood). Large logs (> 26 cm) decayed 40% more slowly than small logs (< 25 cm). The initial volume of logs was 37.6 m3/ha but increased to 44.8 m3/ha after six years. In addition, there was a large shift in the decay-class distribution. The volume of logs in early and late decay classes increased by 71% and 45%, respectively, while the volume of logs in the intermediate decay classes decreased by 32%. The fluctuations appear to result from pulses in mortality, driven by a combination of strong winds and the heart-rot fungus, Phellinus chrysoloma, which was present in more than 30% of all logs at an early stage of decay. These results show that large temporal fluctuations in dead wood also occur in the absence of large-scale disturbance, and that heart-rot fungi are important factors driving the overall dynamics of dead wood. Since many wood-inhabiting species are naturally rare and have very specific substrate demands, such temporal variability in dead wood availability may have effects on biodiversity and should be taken into account when designing small, protected forest areas.