Vegetation-fire feedbacks increase subtropical wildfire risk in scrubland and reduce it in forests.

Climate dictates wildfire activity around the world. But East and Southeast Asia are an apparent exception as fire-activity variation there is unrelated to climatic variables. In subtropical China, fire activity decreased by 80% between 2003 and 2020 amid increased fire risks globally. Here, we assessed the fire regime, vegetation structure, fuel flammability and their interactions across subtropical Hubei, China. We show that tree basal area (TBA) and fuel flammability explained 60% of fire-frequency variance. Fire frequency and fuel flammability, in turn, explained 90% of TBA variance. These results reveal a novel system of scrubland-forest stabilized by vegetation-fire feedbacks. Frequent fires promote the persistence of derelict scrubland through positive vegetation-fire feedbacks; in forest, vegetation-fire feedbacks are negative and suppress fire. Thus, we attribute the decrease in wildfire activity to reforestation programs that concurrently increase forest coverage and foster negative vegetation-fire feedbacks that suppress wildfire.

Large old trees increase growth under shifting climatic constraints: Aligning tree longevity and individual growth dynamics in primary mountain spruce forests.

In a world of accelerating changes in environmental conditions driving tree growth, tradeoffs between tree growth rate and longevity could curtail the abundance of large old trees (LOTs), with potentially dire consequences for biodiversity and carbon storage. However, the influence of tree-level tradeoffs on forest structure at landscape scales will also depend on disturbances, which shape tree size and age distribution, and on whether LOTs can benefit from improved growing conditions due to climate warming. We analyzed temporal and spatial variation in radial growth patterns from ~5000 Norway spruce (Picea abies [L.] H. Karst) live and dead trees from the Western Carpathian primary spruce forest stands. We applied mixed-linear modeling to quantify the importance of LOT growth histories and stand dynamics (i.e., competition and disturbance factors) on lifespan. Finally, we assessed regional synchronization in radial growth variability over the 20th century, and modeled the effects of stand dynamics and climate on LOTs recent growth trends. Tree age varied considerably among forest stands, implying an important role of disturbance as an age constraint. Slow juvenile growth and longer period of suppressed growth prolonged tree lifespan, while increasing disturbance severity and shorter time since last disturbance decreased it. The highest age was not achieved only by trees with continuous slow growth, but those with slow juvenile growth followed by subsequent growth releases. Growth trend analysis demonstrated an increase in absolute growth rates in response to climate warming, with late summer temperatures driving the recent growth trend. Contrary to our expectation that LOTs would eventually exhibit declining growth rates, the oldest LOTs (>400 years) continuously increase growth throughout their lives, indicating a high phenotypic plasticity of LOTs for increasing biomass, and a strong carbon sink role of primary spruce forests under rising temperatures, intensifying droughts, and increasing bark beetle outbreaks.

Factors influencing productivity of pine-dominated stands in South Korea.

Stand productivity research has mainly focused on increasing yield and has recently begun to examine changes in carbon storage. The Korean government is interested in finding ways to increase forest carbon capture to meet carbon neutrality requirements because approximately 63% of the land is covered by forests. In addition, 69% of these forests are older than 30 years old and their productivity and aboveground carbon storage rates are expected to decline. The purpose of this study was to examine the effect of quadratic mean diameter (QMD), stand basal area, site index, slope, climate (MAT and MAP), stand age, stand structural diversity, and stand composition on the productivity of aging Korean red pine-dominated stands. Based on the effects of these factors, we explored how to manage pine forests with the focus of increasing their productivity. Random forest regression was used for the analysis, and periodic basal area increment (PBAI) was used as the dependent variable of stand productivity. Our results show that the most influential factor on stand productivity was QMD. PBAI dramatically decreased from approximately 0.8 to 0.53 m2/ha·year as QMD increased up to 18 cm. Since diameter (QMD) increment is closely associated with changes in tree competition; increasing tree competition with increasing QMD and stand basal area may lead to decreases in PBAI owing to decreases in growth rate due to space and resource limitations and increases in mortality. PBAI decreased when basal area increased from 22 to 51.5 m2/ha. PBAI increased for site index values between 8 and 12.5 m and decreased for stand age values up to approximately 31 years. For climate factors, PBAI generally increased with increasing MAP and slightly increased as MAT increased up to approximately 11.2 °C and then decreased at higher MAT. PBAI initially increased with increasing slope values, decreased with values lower than 15°, and remained stable at slope values in the range of 16-34°. Stand structural diversity, which ranged from 1.32 to 1.62, exhibited a similar negative influence on PBAI associated with increasing stand density. With regard to pine composition, pine stands with a large proportion of pine basal area, showed higher productivity due to the simple stand structure resulting in better growth of shade intolerant pine. This study found that stand density increases with the development of pine stands and that density increases had negative influences on stand productivity. Collectively, our results suggest that stand density management is essential for increasing stand productivity and carbon sequestration in the Korean red pine-dominated stands of South Korea.

Tree growth response to drought partially explains regional-scale growth and mortality patterns in Iberian forests.

Tree-ring data has been widely used to inform about tree growth responses to drought at the individual scale, but less is known about how tree growth sensitivity to drought scales up driving changes in forest dynamics. Here, we related tree-ring growth chronologies and stand-level forest changes in basal area from two independent data sets to test if tree-ring responses to drought match stand forest dynamics (stand basal area growth, ingrowth, and mortality). We assessed if tree growth and changes in forest basal area covary as a function of spatial scale and tree taxa (gymnosperm or angiosperm). To this end, we compared a tree-ring network with stand data from the Spanish National Forest Inventory. We focused on the cumulative impact of drought on tree growth and demography in the period 1981-2005. Drought years were identified by the Standardized Precipitation Evapotranspiration Index, and their impacts on tree growth by quantifying tree-ring width reductions. We hypothesized that forests with greater drought impacts on tree growth will also show reduced stand basal area growth and ingrowth and enhanced mortality. This is expected to occur in forests dominated by gymnosperms on drought-prone regions. Cumulative growth reductions during dry years were higher in forests dominated by gymnosperms and presented a greater magnitude and spatial autocorrelation than for angiosperms. Cumulative drought-induced tree growth reductions and changes in forest basal area were related, but initial stand density and basal area were the main factors driving changes in basal area. In drought-prone gymnosperm forests, we observed that sites with greater growth reductions had lower stand basal area growth and greater mortality. Consequently, stand basal area, forest growth, and ingrowth in regions with large drought impacts was significantly lower than in regions less impacted by drought. Tree growth sensitivity to drought can be used as a predictor of gymnosperm demographic rates in terms of stand basal area growth and ingrowth at regional scales, but further studies may try to disentangle how initial stand density modulates such relationships. Drought-induced growth reductions and their cumulative impacts have strong potential to be used as early-warning indicators of regional forest vulnerability.

Evaluating multiple historical climate products in ecological models under current and projected temperatures.

Gridded historical climate products (GHCPs) are employed with increasing frequency when modeling ecological phenomena across large scales and predicting ecological responses to projected climate changes. Concurrently, there is an increasing acknowledgement of the need to account for uncertainty when employing climate projections from ensembles of global circulation models (GCMs) and emissions scenarios. Despite the growing usage and documented differences among GHCPs, uncertainty characterization has primarily focused on GCM and emissions scenario choice, while the consequences of using a single GHCP to make predictions over space and time have received less attention. Here we employ average July temperature data from observations and seven GHCPs to model plant canopy cover and tree basal area across central Alaska, USA. We first compare the fit of, and support for, models employing observed temperatures, GHCP temperatures, and GHCP temperatures with an elevation adjustment, finding (1) greater support for, and better fit using, elevation-adjusted vs. raw temperature models and (2) overall similar fits of elevation-adjusted models employing temperatures from observations or GHCPs. Focusing on basal area, we next compare predictions generated by elevation-adjusted models employing GHCP data under current conditions and a warming scenario of current temperatures plus 2°C, finding good agreement among GHCPs though with between-GHCP differences and variation primarily at middle elevations (~1,000 m). These differences were amplified under the warming scenario. Finally, using pooled indices of prediction variation and difference across GHCP models, we identify characteristics of areas most likely to exhibit prediction uncertainty under current and warming conditions. Despite (1) overall good performance of GHCP data relative to observations in models and (2) positive correlation among model predictions, variation in predictions across models, particularly in mid-elevation areas where the position of treeline may be changing, suggests researchers should exercise caution if selecting a single GHCP for use in models. We recommend the use of multiple GHCPs to provide additional uncertainty information beyond standard estimated prediction intervals, particularly when model predictions are employed in conservation planning.

Competition indices after reduced impact logging in the Brazilian Amazon.

Competition for resources can affect growth and increase mortality in forest stands. The effects of this process are little known, especially in areas subjected to forest management which show distinct growth dynamics when compared to unmanaged natural areas. This study aimed to estimate, evaluate and select the best fit competition indices (CI) for individual trees in a managed forest in the eastern Amazon. The data used originated from 18 permanent plots of 1 ha each, which were monitored for 12 years after Reduced Impact Logging (RIL). For the competition analysis, 23 indices were tested. CIs were evaluated based on graphical analyses and linear correlation coefficients(r)between each index and variables, basal area growth(ΔG), probability of mortality(Pm), and post-logging period (PLP). In addition, the partial F-test was applied to verify how well the calculated CIs explain the variations observed in ΔGandP(m). The distance-independent index for BAL (Basal Area Larger) performed the best. For the distance-dependent and semi-independent indices, the best interactions with the variables tested occurred in radii of 15 and 20 m away from the object tree. The largest trees presented higher values ofΔG. While P(m)was similar among the three tree classes, larger trees had a slightly higher P(m)due to thier low density of individuals. The results obtained in this study may help to include CIs in growth and production models at the individual tree level in tropical rainforests, especially in those managed in the Brazilian Amazon.

Tropical tree size-frequency distributions from airborne lidar.

In tropical rainforests, tree size and number density are influenced by disturbance history, soil, topography, climate, and biological factors that are difficult to predict without detailed and widespread forest inventory data. Here, we quantify tree size-frequency distributions over an old-growth wet tropical forest at the La Selva Biological Station in Costa Rica by using an individual tree crown (ITC) algorithm on airborne lidar measurements. The ITC provided tree height, crown area, the number of trees >10 m height and, predicted tree diameter, and aboveground biomass from field allometry. The number density showed strong agreement with field observations at the plot- (97.4%; 3% bias) and tree-height-classes level (97.4%; 3% bias). The lidar trees size spectra of tree diameter and height closely follow the distributions measured on the ground but showed less agreement with crown area observations. The model to convert lidar-derived tree height and crown area to tree diameter produced unbiased (0.8%) estimates of plot-level basal area and with low uncertainty (6%). Predictions on basal area for tree height classes were also unbiased (1.3%) but with larger uncertainties (22%). The biomass estimates had no significant bias at the plot- and tree-height-classes level (-5.2% and 2.1%). Our ITC method provides a powerful tool for tree- to landscape-level tropical forest inventory and biomass estimation by overcoming the limitations of lidar area-based approaches that require local calibration using a large number of inventory plots.

Environmental drivers interactively affect individual tree growth across temperate European forests.

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global-change drivers such as climate change or atmospheric deposition, as well as to local land-use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global-change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global-change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global-change drivers, with species-specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus' growth, highlighting species-specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus' growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global-change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.

Fire deficits have increased drought sensitivity in dry conifer forests: Fire frequency and tree-ring carbon isotope evidence from Central Oregon.

A century of fire suppression across the Western United States has led to more crowded forests and increased competition for resources. Studies of forest thinning or stand conditions after mortality events have provided indirect evidence for how competition can promote drought stress and predispose forests to severe fire and/or bark beetle outbreaks. Here, we demonstrate linkages between fire deficits and increasing drought stress through analyses of annually resolved tree-ring growth, fire scars, and carbon isotope discrimination (Δ13 C) across a dry mixed-conifer forest landscape. Fire deficits across the study area have increased the sensitivity of leaf gas exchange to drought stress over the past >100 years. Since 1910, stand basal area in these forests has more than doubled and fire-return intervals have increased from 25 to 140 years. Meanwhile, the portion of interannual variation in tree-ring Δ13 C explained by the Palmer Drought Severity Index has more than doubled in ca. 300-500-year-old Pinus ponderosa as well as in fire-intolerant, ca. 90-190-year-old Abies grandis. Drought stress has increased in stands with a basal area of ≥25 m2 /ha in 1910, as indicated by negative temporal Δ13 C trends, whereas stands with basal area ≤25 m2 /ha in 1910, due to frequent or intense wildfire activity in decades beforehand, were initially buffered from increased drought stress and have benefited more from rising ambient carbon dioxide concentrations, [CO2 ], as demonstrated by positive temporal Δ13 C trends. Furthermore, the average Δ13 C response across all P. ponderosa since 1830 indicates that photosynthetic assimilation rates and stomatal conductance have been reduced by ~10% and ~20%, respectively, compared to expected trends due to increasing [CO2 ]. Although disturbance legacies contribute to local-scale intensity of drought stress, fire deficits have reduced drought resistance of mixed-conifer forests and made them more susceptible to challenges by pests and pathogens and other disturbances.

Asymmetric competition, ontogenetic growth and size inequality drive the difference in productivity between two-strata and one-stratum forest stands.

Size inequality has been considered a key feature of plant population structure with impacts on ecosystem functions. In forest ecosystems, studies examining the relationship between tree size inequality and stand productivity have produced mixed outcomes. These studies found positive, neutral or negative relationships and discussed how this could be influenced by competition for light between trees (e.g. light interception efficiency), but far less attention has been paid to the role played by tree ontogenetic growth. In this article, we present a simple mathematical model that predicts the basal area growth of a two-strata stand as a function of tree basal areas and asymmetric competition. Comparing the growth of this stand to the growth of a spatially homogeneous one-stratum stand and a spatially heterogeneous one-stratum stand, we show that higher growth of the two-strata stand is achieved for concave shape, increasing functions of ontogenetic growth and for low intensities of absolute size-asymmetric competition. We also demonstrate that the difference in growth between the two-strata stand and the one-stratum stands depends on tree size inequality, mean tree basal area and total basal area in the two-strata stand. We finally found that the relationships between tree size inequality and productivity can vary from positive to negative and even non-monotonous. However, we highlight that negative relationships may be more frequent. As a conclusion, our results indicate that ontogenetic growth can have a major impact on the form and the magnitude of the size inequality-productivity relationship.

Size-growth asymmetry is not consistently related to productivity across an eastern US temperate forest network.

Modeling and forecasting forests as carbon sinks require that we understand the primary factors affecting productivity. One factor thought to be positively related to stand productivity is the degree of asymmetry, or the slope of the relationship between tree size and biomass growth. Steeper slopes indicate disproportionate productivity of big trees relative to small trees. Theoretically, big trees outcompete smaller trees during favorable growth conditions because they maintain better access to light. For this reason, high productivity forests are expected to have asymmetric growth. However, empirical studies do not consistently support this expectation, and those that do are limited in spatial or temporal scope. Here, we analyze size-growth relationships from 1970 to 2011 across a diverse network of forest sites in the eastern United States (n = 16) to test whether asymmetry is consistently related to productivity. To investigate this relationship, we analyze asymmetry-productivity relationships between our 16 forests at non-overlapping annual, 2-, 5-, 10-, and 20-year sampling intervals and find that asymmetry is negatively related to productivity, but the strength depends on the specific interval considered. Within-site temporal variability in asymmetry and productivity are generally positively correlated over time, except at the 5-year remeasurement interval. Rather than confirming or failing to support a positive relationship between asymmetry and productivity, our findings suggest caution interpreting these metrics since the relationship varies across forest types and temporal scales.

Increased water-use efficiency translates into contrasting growth patterns of Scots pine and sessile oak at their southern distribution limits.

In forests, the increase in atmospheric CO2 concentrations (Ca ) has been related to enhanced tree growth and intrinsic water-use efficiency (iWUE). However, in drought-prone areas such as the Mediterranean Basin, it is not yet clear to what extent this "fertilizing" effect may compensate for drought-induced growth reduction. We investigated tree growth and physiological responses at five Scots pine (Pinus sylvestris L.) and five sessile oak (Quercus petraea (Matt.) Liebl.) sites located at their southernmost distribution limits in Europe for the period 1960-2012 using annually resolved tree-ring width and δ13 C data to track ecophysiological processes. Results indicated that all 10 natural stands significantly increased their leaf intercellular CO2 concentration (Ci ), and consequently iWUE. Different trends in the theoretical gas-exchange scenarios as a response to increasing Ca were found: generally, Ci tended to increase proportionally to Ca , except for trees at the driest sites in which Ci remained constant. Ci from the oak sites displaying higher water availability tended to increase at a comparable rate to Ca . Multiple linear models fitted at site level to predict basal area increment (BAI) using iWUE and climatic variables better explained tree growth in pines (31.9%-71.4%) than in oak stands (15.8%-46.8%). iWUE was negatively linked to pine growth, whereas its effect on growth of oak differed across sites. Tree growth in the western and central oak stands was negatively related to iWUE, whereas BAI from the easternmost stand was positively associated with iWUE. Thus, some Q. petraea stands might have partially benefited from the "fertilizing" effect of rising Ca , whereas P. sylvestris stands due to their strict closure of stomata did not profit from increased iWUE and consequently showed in general growth reductions across sites. Additionally, the inter-annual variability of BAI and iWUE displayed a geographical polarity in the Mediterranean.

Limited evidence for CO2 -related growth enhancement in northern Rocky Mountain lodgepole pine populations across climate gradients.

Forests sequester large amounts of carbon annually and are integral in buffering against effects of global change. Increasing atmospheric CO2 may enhance photosynthesis and/or decrease stomatal conductance (gs ) thereby enhancing intrinsic water-use efficiency (iWUE), having potential indirect and direct benefits to tree growth. While increasing iWUE has been observed in most trees globally, enhanced growth is not ubiquitous, possibly due to concurrent climatic constraints on growth. To investigate our incomplete understanding of interactions between climate and CO2 and their impacts on tree physiology and growth, we used an environmental gradient approach. We combined dendrochronology with carbon isotope analysis (δ13 C) to assess the covariation of basal area increment (BAI) and iWUE over time in lodgepole pine. Trees were sampled at 18 sites spanning two climatically distinct elevation transects on the lee and windward sides of the Continental Divide, encompassing the majority of lodgepole pine's northern Rocky Mountain elevational range. We analyzed BAI and iWUE from 1950 to 2015, and explored correlations with monthly climate variables. As expected, iWUE increased at all sites. However, concurrent growth trends depended on site climatic water deficit (CWD). Significant growth increases occurred only at the driest sites, where increases in iWUE were strongest, while growth decreases were greatest at sites where CWD has been historically lowest. Late summer drought of the previous year negatively affected growth across sites. These results suggest that increasing iWUE, if strong enough, may indirectly benefit growth at drier sites by effectively extending the growing season via reductions in gs . Strong growth decreases at high elevation windward sites may reflect increasing water stress as a result of decreasing snowpack, which was not offset by greater iWUE. Our results imply that increasing iWUE driven by decreasing gs may benefit tree growth in limited scenarios, having implications for future carbon uptake potential of semiarid ecosystems.

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex.

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40-year tree ring record and a 30-year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (-10 days) due to winter warming and earlier growth cessation (-26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving-window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate-growth correlations matches the start of the recent atmospheric warming pause also known as the 'climate hiatus'. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.

Integrating field and satellite data for spatially explicit inference on the density of threatened arboreal primates.

Spatially explicit models of animal abundance are a critical tool to inform conservation planning and management. However, they require the availability of spatially diffuse environmental predictors of abundance, which may be challenging, especially in complex and heterogeneous habitats. This is particularly the case for tropical mammals, such as nonhuman primates, that depend on multi-layered and species-rich tree canopy coverage, which is usually measured through a limited sample of ground plots. We developed an approach that calibrates remote-sensing imagery to ground measurements of tree density to derive basal area, in turn used as a predictor of primate density based on published models. We applied generalized linear models (GLM) to relate 9.8-ha ground samples of tree basal area to various metrics extracted from Landsat 8 imagery. We tested the potential of this approach for spatial inference of animal density by comparing the density predictions for an endangered colobus monkey, to previous estimates from field transect counts, measured basal area, and other predictors of abundance. The best GLM had high accuracy and showed no significant difference between predicted and observed values of basal area. Our species distribution model yielded predicted primate densities that matched those based on field measurements. Results show the potential of using open-access and global remote-sensing data to derive an important predictor of animal abundance in tropical forests and in turn to make spatially explicit inference on animal density. This approach has important, inherent applications as it greatly magnifies the relevance of abundance modeling for informing conservation. This is especially true for threatened species living in heterogeneous habitats where spatial patterns of abundance, in relation to habitat and/or human disturbance factors, are often complex and, management decisions, such as improving forest protection, may need to be focused on priority areas.

Historical harvests reduce neighboring old-growth basal area across a forest landscape.

While advances in remote sensing have made stand, landscape, and regional assessments of the direct impacts of disturbance on forests quite common, the edge influence of timber harvesting on the structure of neighboring unharvested forests has not been examined extensively. In this study, we examine the impact of historical timber harvests on basal area patterns of neighboring old-growth forests to assess the magnitude and scale of harvest edge influence in a forest landscape of western Oregon, USA. We used lidar data and forest plot measurements to construct 30-m resolution live tree basal area maps in lower and middle elevation mature and old-growth forests. We assessed how edge influence on total, upper canopy, and lower canopy basal area varied across this forest landscape as a function of harvest characteristics (i.e., harvest size and age) and topographic conditions in the unharvested area. Upper canopy, lower canopy, and total basal area increased with distance from harvest edge and elevation. Forests within 75 m of harvest edges (20% of unharvested forests) had 4% to 6% less live tree basal area compared with forest interiors. An interaction between distance from harvest edge and elevation indicated that elevation altered edge influence in this landscape. We observed a positive edge influence at low elevations (<800 m) and a negative edge influence at moderate to high elevations (>800 m). Surprisingly, we found no or weak effects of harvest age (13-60 yr) and harvest area (0.2-110 ha) on surrounding unharvested forest basal area, implying that edge influence was relatively insensitive to the scale of disturbance and multi-decadal recovery processes. Our study indicates that the edge influence of past clearcutting on the structure of neighboring uncut old-growth forests is widespread and persistent. These indirect and diffuse legacies of historical timber harvests complicate forest management decision-making in old-growth forest landscapes by broadening the traditional view of stand boundaries. Furthermore, the consequences of forest harvesting may reach across ownership boundaries, highlighting complex governance issues surrounding landscape management of old-growth forests.

Seasonal patterns of bole water content in old growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco).

Large conifer trees in the Pacific Northwest, USA (PNW) use stored water to extend photosynthesis, both diurnally and seasonally. This is particularly important during the summer drought, which is characteristic of the region. In the PNW, climate change is predicted to result in hotter, drier summers and warmer, wetter winters with decreased snowpack by mid-century. Understanding seasonal bole water dynamics in relation to climate factors will enhance our ability to determine the vulnerability of forests to climate change. Seasonal patterns of bole water content in old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees were studied in the Cascade Mountains of western Oregon, USA. Relative water content (RWC) was monitored hourly in three 400+ and three ~150 years-old trees using permanently mounted dielectric devices for 10 years. RWC increased during the late spring and early summer to maximum levels in August then decreased into fall and remained low over winter. The difference between minimum RWC in the winter and maximum in mid-summer averaged 4.5 and 2.3% for the older and younger trees, respectively, across all years. RWC closely followed growth and was positively correlated with air and soil temperature, vapor pressure deficit and photosynthetically active radiation, but lagged plant available soil water. The progressive decrease in RWC seen each year from mid-summer through fall was attributed to net daily loss of water during the summer drought. The marked increase in RWC observed from spring to mid-summer each year was hypothesized to be the period of embolism repair and water recharge in elastic tissues. We conclude that bole water content is an integral part of tree water dynamics enabling trees to extend carbon assimilation into drought periods and during periods when cold soil inhibits water uptake by roots, an adaptation that could benefit the survival of large PNW trees under climate change.

Basal area growth, carbon isotope discrimination, and intrinsic water use efficiency after fertilization of Douglas-fir in the Oregon Coast Range.

Many hectares of intensively managed Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands in western North America are fertilized with nitrogen (N) to increase growth rates, but only about ⅔ of all stands respond. Understanding the mechanisms of response facilitates prioritization of stands for treatment. The primary objective of this study was to test the hypothesis that the short-term basal area growth response to a single application of 224 kg N ha-1 as urea was associated with reduced stable carbon isotope discrimination (Δ13C) and increased intrinsic water use efficiency (iWUE) in a 20-yr-old plantation of Douglas-fir in the Oregon Coast Range, USA. Increment cores were measured to estimate earlywood, latewood, and total basal area increment over a time series from 1997 to 2015. Stable carbon isotope discrimination and iWUE were estimated using earlywood and latewood stable carbon isotope concentrations in tree-ring holocellulose starting seven years before fertilization in early 2009 and ending seven years after treatment. A highly significant (p<0.01) interaction effect between fertilization treatment and year was found for total basal area growth and earlywood basal area increment. Specifically, fertilized trees showed significant responses (p<0.05) in total basal area growth and earlywood basal area increment in the first (2009) and second (2010) growing seasons after fertilization in 2009. A marginally significant (p<0.10) fertilization effect was found for latewood basal area increment only in the first growing season after treatment. A significant treatment x year interaction was also found for Δ13C and iWUE in earlywood and latewood. Fertilization significantly reduced earlywood Δ13C and increased earlywood iWUE in the first and second growing seasons after fertilization. Only a marginally significant fertilization effect was detected for latewood Δ13C and iWUE in the second growing season after treatment. Previous studies of N fertilization of Douglas-fir forests have reported consistently increased growth and iWUE on low productivity sites treated with relatively high fertilization rates. This study suggested that these responses can also be observed on highly productive sites despite their lower frequency and apparently shorter duration. Other key mechanisms driving growth responses appear less important than iWUE, including an increase in LAI and shift from belowground to aboveground carbon allocation.

Measurements of stem diameter: implications for individual- and stand-level errors.

Stem diameter is one of the most common measurements made to assess the growth of woody vegetation, and the commercial and environmental benefits that it provides (e.g. wood or biomass products, carbon sequestration, landscape remediation). Yet inconsistency in its measurement is a continuing source of error in estimates of stand-scale measures such as basal area, biomass, and volume. Here we assessed errors in stem diameter measurement through repeated measurements of individual trees and shrubs of varying size and form (i.e. single- and multi-stemmed) across a range of contrasting stands, from complex mixed-species plantings to commercial single-species plantations. We compared a standard diameter tape with a Stepped Diameter Gauge (SDG) for time efficiency and measurement error. Measurement errors in diameter were slightly (but significantly) influenced by size and form of the tree or shrub, and stem height at which the measurement was made. Compared to standard tape measurement, the mean systematic error with SDG measurement was only -0.17 cm, but varied between -0.10 and -0.52 cm. Similarly, random error was relatively large, with standard deviations (and percentage coefficients of variation) averaging only 0.36 cm (and 3.8%), but varying between 0.14 and 0.61 cm (and 1.9 and 7.1%). However, at the stand scale, sampling errors (i.e. how well individual trees or shrubs selected for measurement of diameter represented the true stand population in terms of the average and distribution of diameter) generally had at least a tenfold greater influence on random errors in basal area estimates than errors in diameter measurements. This supports the use of diameter measurement tools that have high efficiency, such as the SDG. Use of the SDG almost halved the time required for measurements compared to the diameter tape. Based on these findings, recommendations include the following: (i) use of a tape to maximise accuracy when developing allometric models, or when monitoring relatively small changes in permanent sample plots (e.g. National Forest Inventories), noting that care is required in irregular-shaped, large-single-stemmed individuals, and (ii) use of a SDG to maximise efficiency when using inventory methods to assess basal area, and hence biomass or wood volume, at the stand scale (i.e. in studies of impacts of management or site quality) where there are budgetary constraints, noting the importance of sufficient sample sizes to ensure that the population sampled represents the true population.

Heavy and frequent thinning promotes drought adaptation in Pinus sylvestris forests.

Droughts and their negative effects on forest ecosystems are projected to increase under climate change for many regions. It has been suggested that intensive thinning could reduce drought impacts on established forests in the short-term. Most previous studies on the effect of thinning on drought impacts, however, have been confined to single forest sites. It is therefore still unclear how general and persisting the benefits of thinning are. This study assesses the potential of thinning to increase drought tolerance of the wide spread Scots pine (Pinus sylvestris) in Central Europe. We hypothesized (1) that increasing thinning intensity benefits the maintenance of radial growth of crop trees during drought (resistance) and its recovery following drought, (2) that those benefits to growth decrease with time elapsed since the last thinning and with stand age, and (3) that they may depend on drought severity as well as water limitations in pre- and post-drought periods. To test these hypotheses, we assessed the effects of thinning regime, stand age, and drought severity on radial growth of 129 Scots pine trees during and after drought events in four long-term thinning experiments in Germany. We found that thinning improved the recovery of radial growth following drought and to a lesser extent the growth resistance during a drought event. Growth recovery following drought was highest after the first thinning intervention and in recently and heavily thinned stands. With time since the last thinning, however, this effect decreased and could even become negative when compared to unthinned stands. Further, thinning helped to avoid an age-related decline in growth resistance (and recovery) following drought. The recovery following drought, but not the resistance during drought, was related to water limitations in the drought period. This is the first study that analyzed drought-related radial growth in trees of one species across several stands of different age. The interaction between thinning intensity and time since the last thinning underline the importance to distinguish between short- and long-term effects of thinning. According to our analysis, only thinning regimes, with relatively heavy and frequent thinning interventions would increase drought tolerance in pine stands.