Ozone uptake, water loss and carbon exchange dynamics in annually drought-stressed Pinus ponderosa forests: measured trends and parameters for uptake modeling

This paper describes 3 years of physiological measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing along an ozone concentration gradient in the Sierra Nevada, California, including variables necessary to parameterize, validate and modify photosynthesis and stomatal conductance algorithms used to estimate ozone uptake. At all sites, gas exchange was under tight stomatal control during the growing season. Stomatal conductance was strongly correlated with leaf water potential (R2=0.82), which decreased over the growing season with decreasing soil water content (R2=0.60). Ozone uptake, carbon uptake, and transpirational water loss closely followed the dynamics of stomatal conductance. Peak ozone and CO2 uptake occurred in early summer and declined progressively thereafter. As a result, periods of maximum ozone uptake did not correspond to periods of peak ozone concentration, underscoring the inappropriateness of using current metrics based on concentration (e.g., SUM0, W126 and AOT40) for assessing ozone exposure risk to plants in this climate region. Both Jmax (maximum CO2-saturated photosynthetic rate, limited by electron transport) and Vcmax (maximum rate of Rubisco-limited carboxylation) increased toward the middle of the growing season, then decreased in September. Intrinsic water-use efficiency rose with increasing drought stress, as expected. The ratio of Jmax to Vcmax was similar to literature values of 2.0. Nighttime respiration followed a Q10 of 2.0, but was significantly higher at the high-ozone site. Respiration rates decreased by the end of the summer as a result of decreased metabolic activity and carbon stores.     

Tree mortality in drought-stressed mixed-conifer and ponderosa pine forests, Arizona, USA

We monitored tree mortality in northern Arizona (USA) mixed-conifer and ponderosa pine (Pinus ponderosa Dougl. ex Laws) forests from 1997 to 2007, a period of severe drought in this area. Mortality was pervasive, occurring on 100 and 98% of 53 mixed-conifer and 60 ponderosa pine plots (1-ha each), respectively. Most mortality was attributable to a suite of forest insects, mediated by drought stress. The number of trees dying from 2002 to 2007 was more than 200% greater than the number dying from 1997 to 2002 in mixed-conifer forest and 74% greater in ponderosa pine forest. Extent of mortality was spatially variable in both forest types. Median cumulative mortality (the ratio of dead to live trees) increased by approximately 53 and 65% in mixed-conifer and ponderosa pine forests, respectively, from 2002 to 2007. Median mortality rates from 2002 to 2007 were approximately 2.0% year⁻¹ in mixed-conifer forest (range = 0-28.5%) and 0.4% year⁻¹ in ponderosa pine forest (range = 0-13.6%). Mortality rates generally were not strongly related to either elevation or stand density. Mortality was nonrandom with respect to tree size classes and species. Proportions of trees dying were greatest in the largest size classes, particularly in mixed-conifer forest, where mortality in the largest size class exceeded 22% from 2002 to 2007. Mortality in mixed-conifer forest was particularly pronounced for quaking aspen (85%) and white fir (28%), the least drought tolerant species present. These results provide an early glimpse of how these forest types are likely to respond to predicted climate changes in the southwestern USA. They suggest that these forests are not resilient to climate change, and that treatments to increase resilience to climate change may be appropriate. Research on causes of spatial heterogeneity in extent of mortality might suggest valuable approaches to aid in increasing resilience.     

Accentuation of gas exchange gradients in flushes of ponderosa pine exposed to ozone

Two-year-old ponderosa pine seedlings (Pinus ponderosa Laws.) were exposed to episodic O(3) concentrations in open-top chambers for two consecutive growing seasons (June through September of 1990 and 1991). Near the end of the second season of O(3) exposure, gas exchange was measured on needles of surviving flushes at saturating CO(2) and photosynthetic photon flux density (PPFD). Both photosynthetic capacity (A(sat)) and stomatal conductance to water vapor (g(wv)) declined linearly with needle age but differences within a flush were also found. Gas exchange rates of needles from the base of the current-year flush were significantly lower than those of needles from the top of the flush, even though age differences between needles were negligible. Although most measurements were conducted at saturating CO(2), similar patterns of gas exchange were also found at 350 micro mol mol(-1) CO(2), indicating that photosynthesis of needles at the bottom of the flush was more strongly affected by O(3) than that of needles at the top of the flush, even though the potential for O(3) uptake was probably less in needles at the bottom of the flush because of reduced stomatal conductance. Carboxylation efficiency (deltaA/deltaC(i)), the linear slope of the A/C(i) response, was highly correlated with A(sat), varying with needle age, needle position in the flush and O(3) exposure, but the magnitude of the reductions was greater than for A(sat). We conclude that susceptibility to O(3) damage among needles of an individual seedling varies not only with needle age but also with needle position, and that reductions in photosynthetic capacity may not be directly attributable to increased uptake of the pollutant. The data also indicate the need to consider within-flush variation when estimating whole-plant carbon gain and O(3) uptake.     

Pricing ecosystem resilience in frequent-fire ponderosa pine forests

Dry forests across the United States have become subject to declining resilience and, consequently, increased vulnerability to catastrophic wildfires. These disturbances cause severe environmental and social damages and may dislodge the forest into a different ecological regime. Forests provide many valuable services, such as the provisioning of timber and the sequestering of carbon that would otherwise contribute to climate change. The high-severity conflagrations that have become regular occurrences in many dry forests impinge the delivery of such benefits, particularly in the event of a regime shift. Sustainable forest management should take these risks into account. This article analyzes the economics of resilience in dry forests with respect to catastrophic fires and ecological thresholds. We illustrate how to price ecosystem resilience for the fire-prone ponderosa pine forests of the western United States. This analysis demonstrates that pricing forest resilience also establishes the economic value of ecological restoration with respect to ecosystem services, thereby operationalizing forest management as an investment in natural capital.     

Carbon isotopic composition, gas exchange, and growth of three populations of ponderosa pine differing in drought tolerance

Effects of water supply on gas exchange, carbon isotopic composition, and relative growth rate were compared among seedlings from three populations of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) grown in a controlled environment chamber. The three populations were chosen to represent high, moderate and low drought tolerance. There was no indication that drought tolerance was related to high water-use efficiency. Populations differed (P < 0.05) in relative growth rate (RGR), but did not differ (P > 0.10) in gas exchange variables or carbon isotope ratio (delta(13)C). Well-watered seedlings had significantly higher RGR, xylem pressure potential (Psi(xpp)), net photosynthesis (A), stomatal conductance to water vapor (g), and lower delta(13)C and instantaneous water-use efficiency than water-stressed seedlings. With decreasing Psi(xpp), A decreased linearly, whereas g decreased exponentially. Seedlings of the highly drought-tolerant population were more sensitive to water availability than seedlings from the other populations; they used water quickly when water was available, but closed their stomata in response to water stress. We conclude that, in ponderosa pine, the drought avoidance mechanism is more important for survival and growth in arid and semiarid environments than the efficient use of water.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Water limitations to carbon exchange in old-growth and young ponderosa pine stands

We investigated the impact of seasonal soil water deficit on the processes driving net ecosystem exchange of carbon (NEE) in old-growth and recently regenerating ponderosa pine (Pinus ponderosa Doug. ex Laws.) stands in Oregon. We measured seasonal patterns of transpiration, canopy conductance and NEE, as well as soil water, soil temperature and soil respiration. The old-growth stand (O) included two primary age classes (50 and 250 years), had a leaf area index (LAI) of 2.1 and had never been logged. The recently regenerating stand (Y) consisted predominantly of 14-year-old ponderosa pine with an LAI of 1.0. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. By August, soil volumetric water content within the upper 30 cm had declined to a seasonal minimum of 0.07 at both sites. Between April and June, both stands showed similar rates of transpiration peaking at 0.96 mm day(-1); thereafter, trees at the Y site showed increasing drought stress with canopy stomatal resistance increasing 6-fold by mid-August relative to values for trees at the O site. Over the same period, predawn water potential (psi(pd)) of trees at the Y site declined from -0.54 to -1.24 MPa, whereas psi(pd) of trees at the O site remained greater than -0.8 MPa throughout the season. Soil respiration at the O site showed a strong seasonal correlation with soil temperature with no discernible constraints imposed by declining soil water. In contrast, soil respiration at the Y site peaked before seasonal maximal soil temperatures and declined thereafter with declining soil water. No pronounced seasonal pattern in daytime NEE was observed at either site between April and September. At the Y site this behavior was driven by concurrent soil water limitations on soil respiration and assimilation, whereas there was no evidence of seasonal soil water limitations on either process at the O site.     

Response of antelope bitterbrush to repeated prescribed burning in Central Oregon ponderosa pine forests

Antelope bitterbrush is a dominant shrub in many interior ponderosa pine forests in the western United States. How it responds to prescribed fire is not well understood, yet is of considerable concern to wildlife and fire managers alike given its importance as a browse species and as a ladder fuel in these fire-prone forests. We quantified bitterbrush cover, density, and biomass in response to repeated burning in thinned ponderosa pine forests. Low- to moderate-intensity spring burning killed the majority of bitterbrush plants on replicate plots. Moderately rapid recovery of bitterbrush density and cover resulted from seedling recruitment plus limited basal sprouting. Repeated burning after 11 years impeded the recovery of the bitterbrush community. Post-fire seed germination following the repeated burns was 3–14-fold lower compared to the germination rate after the initial burns, while basal sprouting remained fairly minor. After 15 years, bitterbrush cover was 75–92% lower on repeated-burned compared to unburned plots. Only where localized tree mortality resulted in an open stand was bitterbrush recovery robust. By controlling bitterbrush abundance, repeated burning eliminated the potential for wildfire spread when simulated using a customized fire behavior model. The results suggest that repeated burning is a successful method to reduce the long-term fire risk imposed by bitterbrush as an understory ladder fuel in thinned pine stands. Balancing the need to limit fire risk yet provide adequate bitterbrush habitat for wildlife browse will likely require a mosaic pattern of burning at the landscape scale or a burning frequency well beyond 11 years to allow a bitterbrush seed crop to develop.     

RESEARCH ON REGENERATION APPROACHES OF KOREAN PINE FORESTS

Broad-lcavedKoreanpincforcstisthcclimaticclimaxconinunitVofnortheastmoun-tainousregionsofChina.AsthemaintreesPecies,Koreanpinehasexccllentwood.AgreatattentionbasbccnpaidtorecoveringtheseforesttyPesafterharvesting.Additional1y,KoreanpineisoneofkeyregcncrationtreesPeCiesinthisregion.TherehasbeenlargeareaofKorcanpincplantations.Howevcr,thercaresomeproblemsncedingtobestUdicdfwtheramongwhichoneistoselectsuitabIeropenerationapproacl1esofKoreanpincforesls.WHATISTHEPROBLEM?ThcnatUralregen…     

Guidelines for Managing Wildlife Habitats in Southwestern Ponderosa Pine Forests of the United States

IntroductionEcoledcalrelahonshipsanddePendenceofwildlifesPe-ciesonsouthwesternPOnderosapine(Pineponderosa)fOrestsarecloselyrelat6dtohabitatuse.Foraginghabits,seasOndfoodPreferences,anduseoftreesforfeedingarelawtydePendentuponthestrUctureandspahalar-rangme…     

Carbon and nitrogen cycling immediately following bark beetle outbreaks in southwestern ponderosa pine forests

Bark beetle infestation is a well-known cause of historical low-level disturbance in southwestern ponderosa pine forests, but recent fire exclusion and increased tree densities have enabled large-scale bark beetle outbreaks with unknown consequences for ecosystem function. Uninfested and beetle-infested plots (n = 10 pairs of plots on two aspects) of ponderosa pine were compared over one growing season in the Sierra Ancha Experimental Forest, AZ to determine whether infestation was correlated with differences in carbon (C) and nitrogen (N) pools and fluxes in aboveground biomass and soils. Infested plots had at least 80% of the overstory ponderosa pine trees attacked by bark beetles within 2 years of our measurements. Both uninfested and infested plots stored ∼9 kg C m⁻² in aboveground tree biomass, but infested plots held 60% of this aboveground tree biomass in dead trees, compared to 5% in uninfested plots. We hypothesized that decreased belowground C allocation following beetle-induced tree mortality would alter soil respiration rates, but this hypothesis was not supported; throughout the growing season, soil respiration in infested plots was similar to uninfested plots. In contrast, several results supported the hypothesis that premature needlefall from infested trees provided a pulse of low C:N needlefall that altered soil N cycling. The C:N mass ratio of pine needlefall in infested plots (∼45) was lower than uninfested plots (∼95) throughout the growing season. Mineral soils from infested plots had greater laboratory net nitrification rates and field resin bag ammonium accumulation than uninfested plots. As bark beetle outbreaks become increasingly prevalent in western landscapes, longer-term biogeochemical studies on interactions with other disturbances (e.g. fire, harvesting, etc.) will be required to predict changes in ecosystem structure and function.     

Biomass and carbon sequestration of ponderosa pine plantations and native cypress forests in northwest Patagonia

Fast growth tree plantations and secondary forests are considered highly efficient carbon sinks. In northwest Patagonia, more than 2 million ha of rangelands are suitable for forestry, and tree plantation or native forest restoration could largely contribute to climate change mitigation. The commonest baseline is the heavily grazed gramineous steppe of Festuca pallescens (St. Yves) Parodi. To assess the carbon sequestration potential of ponderosa pine (Pinus ponderosa (Dougl.) Laws) plantations and native cypress (Austrocedrus chilensis (Don) Flor. et Boutl.), individual above and below ground biomass models were developed, and scaled to stand level in forests between 600 and 1500 annual rainfall. To calculate the carbon sequestration baseline, the pasture biomass was simulated. Also, soil carbon at two depths was assessed in paired pine-cypress-pasture sample plots, the same as the litter carbon content of both forest types. Individual stem, foliage, branch and root log linear equations adjusted for pine and cypress trees presented similar slopes (P>0.05), although some differed in the elevations. Biomass carbon was 52.3 Mg ha⁻¹ (S.D.=30.6) for pine stands and 73.2 Mg ha⁻¹ (S.D.=95.4) for cypress forests, given stand volumes of 148.1 and 168.4 m³ ha⁻¹, respectively. Soil carbon (litter included) was 86.3 Mg ha⁻¹ (S.D.=46.5) for pine stands and 116.5 Mg ha⁻¹ (S.D.=38.5) for cypress. Root/shoot ratio was 19.5 and 11.4%, respectively. The low r/s value for cypress may account for differences in nutrient cycling and water uptake potential. At stand level, differences in foliage, taproot and soil carbon compartments were highly significative (P<0.01) between both forest types. In pine stands, both biomass and soil carbon were highly explained by the rainfall gradient (r²=0.94). Nevertheless, such a relationship was not found for cypress, possibly due to stand and soil disturbances in sample plots. The carbon baseline estimated in pasture biomass, including litter, was 2.6 Mg ha⁻¹ (S.D.=0.8). Since no differences in soil carbon were found between pasture and both forest types, additionality should be accounted only by biomass. However, the replacement of pasture by pine plantations may decrease the soil carbon storage, at least during the first years. On the other hand, the soil may be a more relevant compartment of sequestered carbon in cypress forests, and if pine plantation replaces cypress forests, soil carbon losses could cause a negative balance.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

Introduction of ponderosa pine and Douglas-fir to Argentina

Patterns of shoot elongation of 2-yr seedlings from native North American populations of ponderosa pine and Douglas-fir were compared to those of Argentine land races originating from unknown provenances. The comparisons were conducted in Moscow, Idaho (USA), and suggested that the ponderosa pine land race was descended from a California provenance at low or middle elevations but that the growth potential of the land race was only mediocre in comparison to eight native populations. The Douglas-fir land race exhibited a relatively high growth potential in comparison to 19 native provenances and undoubtedly originated from a mild coastal environment. The results provide concrete recommendations for upgrading the growth potential of the Argentine land races by importing germ plasm of specific provenances.     

Changes in understory production for three prescribed burns of different ages in ponderosa pine

The general lack of resource response information severely limits economic evaluation of prescribed burning in most forest types. This paper present changes in understory production at three sites on basalt soils following prescribed burning in ponderosa pine (Pinus ponderosa Laws.) in Arizona. The sites were burned during the fall 2, 5, and 7 years before sampling in 1981. Regression equations were developed to predict production from plant basal area for four common grass species; production of remaining herbaceous vegetation was measured by harvesting. Based on the results of our study and other studies of fall prescribed burning on volcanic soils in Arizona ponderosa pine, understory production response appears to be variable for 1–2 years following burning. Herbage production exhibits no change or an increase; forage production exhibits no change or a decrease. Studies of understory response more than 2 years following both wildfire and prescribed burning in Arizona ponderosa pine, however, show a general trend toward increased production. Thus, prescribed burning in southwestern pine on basalt soils produces long-term benefits in increased understory production, particularly in pole stands, which dominate much of the region.     

一次飑线对阔叶红松林危害的记录

红松(Pinus koraiensis)是我国东北林区的珍贵树种,2002年被正式列为黑龙江省二级保护植物,它的优异材质和独特的价值被世界所公认。红松是山地树种,在世界上分布不广,我国东北是其自然分布区的中心地带。阔叶红松林是小兴安岭南坡的地带性植被,是伊春林区的主要森林类型。2005     

Spatially modeling wildland fire severity in pine forests of Galicia,Spain

Fire is a major disturbance in forests and one of the most important carbon emissions sources, which contributes to climate change. Carbon emissions are directly correlated with the degree of organic matter consumption or fire severity. Gaining knowledge about the relative strength of the various explanatory variables is essential to mitigate its environmental impact. We tested an approach that combines wind modeling, light detection and ranging (LiDAR), remotely sensed vegetation indices and topography data for assessing the occurrence of high-severity fire using the random forests ensemble learning method. Data from four wildfires that occurred in Galicia (northwestern Spain) were used to exemplify the application of this approach. The models predicted high-severity occurrence with a classification accuracy ranging from 77 to 94%. High-severity fire occurred more frequently in areas of high simulated wind speed, and more pronouncedly, for cases reported as wind-driven fires. High severity also occurred more frequently in areas of high terrain roughness, on sunny slopes and in low canopy base height stands. This approach allowed predicting spatially explicit fire severity at a mean scale level (resolution of 25 m) with accuracy rates from 80 to 95%. This approach may be helpful for fire managers when delimiting and planning fuel treatments for severity mitigation or during fire suppression, and for post hoc case studies.     

Real versus perceived conflicts between restoration of ponderosa pine forests and conservation of the Mexican spotted owl

Progress in implementing ecosystem approaches to conservation and restoration is slowed by legitimate concerns about the effects of such approaches on individual imperiled species. The perceived conflict between the restoration of fire-excluded forests and concomitant reduction of dense fuels and high-severity wildfire, versus the recovery of endangered species, has led to a policy ambiguity that has slowed on-the-ground action at a time when active management is urgently needed, both for ecosystem restoration and species conservation. The Mexican spotted owl (Strix occidentalis lucida) in the southwestern U.S.A. is emblematic of this perceived conflict, with numerous appeals and lawsuits focused on the species and vast acres of forest managed with habitat quality for this species in mind. We use spatial analysis across large landscapes in Arizona to examine potential conflicts between the desire to reduce the likelihood of uncharacteristically severe wildfire and restore native fire regimes, and the concurrent desire and legal mandate to manage forests for the recovery of the owl. Our spatially explicit analysis indicates that real conflicts between these management objectives exist, but that locations where conflicts might inhibit active forest management represent less than 1/3 of the 811,000 ha study region. Furthermore, within the areas where conflicts might be expected, the majority of the forest could be managed in ways that would reduce fire hazard without eliminating owl habitat. Finally, management treatments that emphasize ecosystem restoration might improve the suitability of large areas of forest habitat in the southwest that is currently unsuitable for owls. These results demonstrate that even where policy conflicts exist, their magnitude has been overstated. Active restoration of dry forests from which fire has been excluded is compatible in many areas with conservation and recovery of the owl. Identifying and prioritizing areas to meet the dual goals of ecosystem restoration and imperiled species conservation require a broad spatial approach that is analytically feasible but currently underutilized. Working together, conservation biologists, restoration ecologists, and forest managers can employ landscape-level spatial analysis to identify appropriate areas for management attention, identify suitable management practices, and explore the predicted consequences of alternative management scenarios on forests, fire ecology, and the fate of sensitive species of conservation concern.     

Assessing and modeling total height and diameter increment of ponderosa pine planted in Minnesota,USA

New Forests - Forest managers are increasingly planting non-native tree species that are adapted to anticipated future conditions such as increased droughts. This work quantified individual tree...