Historical Stem‐Mapped Permanent Plots Increase Precision of Reconstructed Reference Data in Ponderosa Pine Forests of Northern Arizona

Forest structural reference conditions are widely used to understand how ecosystems have been altered and guide restoration and management objectives. We used six stem‐mapped permanent plots established in the early twentieth century to provide precise structural reference conditions for ponderosa pine forests of northern Arizona prior to Euro‐American settlement. Reference conditions for these plots in 1873–1874 included the following historical attributes: tree densities of 45–127 trees/ha, mean tree diameter at breast height (dbh) of 43.8 cm with a corresponding quadratic mean diameter range of 41.5–51.3 cm, and a stand basal area of 9.2–18.0 m²/ha. The reconstructed diameter distributions (for live ponderosa pine trees with dbh ≥9.14 cm) prior to fire exclusion varied in shape but generally displayed an irregular unimodal distribution. We suggest that management objectives for the structural restoration of ponderosa pine forests of northern Arizona emphasize: (1) conservation and retention of all pre‐settlement (>130 years) trees; (2) reduction of tree densities with a restoration objective ranging between 50 and 150 trees/ha having a large‐tree component between 25 and 50% of the total trees per hectare, respectively; (3) manipulation of the diameter distribution to achieve a unimodal or irregular, uneven‐aged shape (possibly targeting a balanced, uneven‐aged shape on cinder soil types) through the use of harvest and thinning practices that mimic gap disturbances (i.e., individual tree selection system); and (4) retention of 3–11 snags and logs per hectare resulting from natural mortality.     

Fire, fuels and restoration of ponderosa pine–Douglas fir forests in the Rocky Mountains, USA

Aim Forest restoration in ponderosa pine and mixed ponderosa pine–Douglas fir forests in the US Rocky Mountains has been highly influenced by a historical model of frequent, low‐severity surface fires developed for the ponderosa pine forests of the Southwestern USA. A restoration model, based on this low‐severity fire model, focuses on thinning and prescribed burning to restore historical forest structure. However, in the US Rocky Mountains, research on fire history and forest structure, and early historical reports, suggest the low‐severity model may only apply in limited geographical areas. The aim of this article is to elaborate a new variable‐severity fire model and evaluate the applicability of this model, along with the low‐severity model, for the ponderosa pine–Douglas fir forests of the Rocky Mountains. Location Rocky Mountains, USA. Methods The geographical applicability of the two fire models is evaluated using historical records, fire histories and forest age‐structure analyses. Results Historical sources and tree‐ring reconstructions document that, near or before ad 1900, the low‐severity model may apply in dry, low‐elevation settings, but that fires naturally varied in severity in most of these forests. Low‐severity fires were common, but high‐severity fires also burned thousands of hectares. Tree regeneration increased after these high‐severity fires, and often attained densities much greater than those reconstructed for Southwestern ponderosa pine forests. Main conclusions Exclusion of fire has not clearly and uniformly increased fuels or shifted the fire type from low‐ to high‐severity fires. However, logging and livestock grazing have increased tree densities and risk of high‐severity fires in some areas. Restoration is likely to be most effective which seeks to (1) restore variability of fire, (2) reverse changes brought about by livestock grazing and logging, and (3) modify these land uses so that degradation is not repeated.     

Stand-replacing wildfires increase nitrification for decades in southwestern ponderosa pine forests

Stand-replacing wildfires are a novel disturbance within ponderosa pine (Pinus ponderosa) forests of the southwestern United States, and they can convert forests to grasslands or shrublands for decades. While most research shows that soil inorganic N pools and fluxes return to pre-fire levels within a few years, we wondered if vegetation conversion (ponderosa pine to bunchgrass) following stand-replacing fires might be accompanied by a long-term shift in N cycling processes. Using a 34-year stand-replacing wildfire chronosequence with paired, adjacent unburned patches, we examined the long-term dynamics of net and gross nitrogen (N) transformations. We hypothesized that N availability in burned patches would become more similar to those in unburned patches over time after fire as these areas become re-vegetated. Burned patches had higher net and gross nitrification rates than unburned patches (P < 0.01 for both), and nitrification accounted for a greater proportion of N mineralization in burned patches for both net (P < 0.01) and gross (P < 0.04) N transformation measurements. However, trends with time-after-fire were not observed for any other variables. Our findings contrast with previous work, which suggested that high nitrification rates are a short-term response to disturbance. Furthermore, high nitrification rates at our site were not simply correlated with the presence of herbaceous vegetation. Instead, we suggest that stand-replacing wildfire triggers a shift in N cycling that is maintained for at least three decades by various factors, including a shift from a woody to an herbaceous ecosystem and the presence of fire-deposited charcoal.     

Recommendations for Integrating Restoration Ecology and Conservation Biology in Ponderosa Pine Forests of the Southwestern United States

Over the past century, ponderosa pine–dominated landscapes of the southwestern United States have been altered by human activities such as grazing, timber harvest, road building, and fire exclusion. Most forested areas within these landscapes now show increased susceptibility to stand‐replacing fires, insect outbreaks, and drought‐related mortality. Recent large wildfires in the region have spurred public interest in large‐scale fuel reduction and restoration programs, which create perceived and real conflicts with the conservation of biodiversity. Conservation concerns include the potential for larger road networks, soil and understory disturbance, exotic plant invasion, and the removal of large trees in treated areas. Pursuing prescribed burning, thinning, or other treatments on the broad scale that many scientists and managers envision requires the reconciliation of ecological restoration with biodiversity conservation. This study presents recommendations from a workshop for integrating the principles and practices of restoration ecology and conservation biology, toward the objective of restoring the composition, structure, and function of dry ponderosa pine forests. Planning on the scale of hundreds of thousands of hectares offers opportunities to achieve multiple objectives (e.g., rare species protection and restoration of ecological structures and processes) that cannot easily be addressed on a site‐by‐site basis. However, restoration must be coordinated with conservation planning to achieve mutual objectives and should include strict guidelines for protection of rare, declining, and sensitive habitats and species.     

Ecosystem Carbon Remains Low for Three Decades Following Fire and Constrains Soil CO2 Responses to Precipitation in Southwestern Ponderosa Pine Forests

The fire regime of ponderosa pine forests in the southwestern United States has shifted over the past century from historically frequent, low-intensity surface fires to infrequent, stand-replacing crown fires. We quantified plant and soil carbon (C) responses to this new fire regime and assessed interactions between changes in fire regime and changes in precipitation regime predicted by some climate models (specifically, an earlier monsoon rain season). We hypothesized that soil C pools and carbon dioxide (CO₂) efflux rates would decrease initially following stand-replacing fires (due to low plant C inputs and the loss of the soil surficial organic (O) horizon), but then increase with time-after-fire (as plant C inputs increase). Water availability often limits soil biological activity in these forests, but we predicted that low soil C availability following fire would constrain soil CO₂ efflux responses to precipitation. In a series of sites with histories of stand-replacing fires that burned between 2 and 34?years prior to sampling, burned patches had lower soil C pools and fluxes than adjacent unburned patches, but there was no evidence of a trend with time-after-fire. Burned forests had 7,500?g C m^?2 less live plant biomass C (P?<?0.001), 1,600?g C m^?2 less soil total C (P?<?0.001) and 90?g C m^?2 less soil labile C (P?<?0.001) than unburned forests. Lower soil labile C in burned patches was due to both a loss of O horizon mass with fire and lower labile C concentrations (g labile C kg^?1 soil total C) in the mineral soil. During the annual drought that precedes summer monsoon rains, both burned and unburned patches had soil CO₂ efflux rates ranging from 0.9 to 1.1?g CO₂-C m^?2 day^?1. During the monsoon season, soil CO₂ efflux in unburned patches increased to approximately 4.8?g CO₂-C m^?2 day^?1 and rates in paired burned patches (3.4?g CO₂-C m^?2 day^?1) were lower (P?<?0.001). We also used field irrigation to experimentally create an earlier and longer monsoon season, and soil CO₂ efflux rates at both burned and unburned plots increased initially in response to watering, but decreased to below control (plots without irrigation) rates within weeks. Watering did not significantly change cumulative growing season soil CO₂ efflux, supporting our prediction that C availability constrains soil CO₂ efflux responses to precipitation. This research advances our understanding of interactions among climate, fire, and C in southwestern forests, suggesting that climate-induced shifts toward more stand-replacing fires will decrease soil C for decades, such that a single fire can constrain future soil biological responses to precipitation regime changes.     

Slash Pile Burning Effects on Soil Biotic and Chemical Properties and Plant Establishment: Recommendations for Amelioration

Ponderosa pine forest restoration consists of thinning trees and reintroducing prescribed fire to reduce unnaturally high tree densities and fuel loads to restore ecosystem structure and function. A current issue in ponderosa pine restoration is what to do with the large quantity of slash that is created from thinning dense forest stands. Slash piling burning is currently the preferred method of slash removal because it allows land managers to burn large quantities of slash in a more controlled environment in comparison with broadcast burning slash. However burning slash piles is known to have adverse effects such as soil sterilization and exotic species establishment. This study investigated the effects of slash pile burning on soil biotic and chemical variables and early herbaceous succession on burned slash pile areas. Slash piles were created following tree thinning in two adjacent approximately 20‐ha ponderosa pine (Pinus ponderosa) restoration treatments in the Coconino National Forest near Flagstaff, Arizona. We selected 30 burned slash pile areas and sampled across a gradient of the burned piles for arbuscular mycorrhizal (AM) propagule densities, the soil seed bank, and soil chemical properties. In addition, we established five 1‐m² plots in each burned pile to quantify the effect of living soil (AM inoculum) and seeding amendments on early herbaceous succession in burned slash pile areas. The five treatments consisted of a control (no treatment), living soil (AM inoculum) amendment, sterilized soil (no AM inoculum) amendment, seed amendment, and a seed/soil (AM inoculum) amendment. Slash pile burning nearly eliminated populations of viable seeds and AM propagules and altered soil chemical properties. Amending scars with native seeds increased the cover of native forbs and grasses. Furthermore adding both seed and living soil more than doubled total native plant cover and decreased ruderal and exotic plant cover. These results indicate that seed/soil amendments that increase native forbs and grasses may enhance the rate of succession in burned slash pile areas by allowing these species to outcompete exotic and ruderal species also establishing at the site through natural regeneration.     

Effects of Restoration Thinning on Presettlement Pinus ponderosa in Northern Arizona

In the 100 years following the arrival of Euro-American settlers in northern Arizona, Pinus ponderosa (ponderosa pine) forests changed from open, low-density stands to closed, high-density stands. The increase in tree density has been detrimental to the vigor of old-growth trees that established before settlement (presettlement trees). In this study, we examined whether the vigor of presettlement trees could be improved by restoring the original stand structure by thinning the ponderosa pines that established after settlement (postsettlement trees). The restoration treatment caused the following changes in the presettlement trees and their environment in the first year following thinning: an increase in volumetric soil water content between May and August, an increase in predawn xylem water potential in July and August, a decrease in midday xylem water potential in June and August, an increase in net photosynthetic rate in August, an increase in foliar nitrogen concentration in July and August, and an increase in bud and needle size. The results show that the thinning restoration treatment improved the condition of presettlement ponderosa pines by increasing canopy growth and the uptake of water, nitrogen, and carbon.     

Fuel-Reduction Treatment Effects on Avian Community Structure and Diversity

Abstract: We assessed responses of the breeding bird community to mechanical thinning and prescribed surface fire, alone and in combination, between 2000 and 2006 in ponderosa pine (Pinus ponderosa) forests in northern Arizona, USA. Fuel-reduction treatments did not affect species richness or evenness, and effects on density of 5 commonly detected species varied among species. Populations of some species, such as the western bluebird (Sialia mexicana), increased following burning treatments, whereas others, such as the mountain chickadee (Poecile gambeli), decreased in response to thinning treatments. Our results also identified a temporal response component, where avian community composition and structure changed synchronously on all treatments over time. Given the modest effects these small-scale fuel-reduction treatments had on avian composition and the specific density responses of particular species, our results suggest that land managers should consider implementing prescribed surface fire after thinning projects, where appropriate.     

Seven-Year Results of Thinning and Burning Restoration Treatments on Old Ponderosa Pines at the Gus Pearson Natural Area

Abstract We examined the 7‐year effects of three restoration treatments on leaf physiology and insect‐resistance characteristics of pre‐settlement age ponderosa pines (Pinus ponderosa Dougl. ex Laws.) at the Gus Pearson Natural Area (GPNA) in northern Arizona. Restoration treatments were: (1) thinned in 1993 to approximate pre‐Euro‐American settlement stand structure, (2) thinned plus prescribed burned in 1994 and 1998, and (3) untreated control. Tree physiological and insect‐resistance characteristics were measured in year 2000, 7 years after thinning, using the same procedures as an earlier study performed in 1996. Consistent with the 1996 results, pre‐dawn water potential in 2000 was consistently lower in the control than both thinned treatments. Both thinned treatments continued to have increased foliar nitrogen concentration in leaves 7 years after treatment. However lower leaf nitrogen concentration in the thinned and burned compared with the thinned treatment suggests lower nitrogen availability to trees in repeatedly burned plots. Analysis of leaf gas exchange characteristics and carbon isotope content (δ¹³C) suggests continued stimulation of photosynthesis by both thinning treatments. Differences among treatments in resin volume, a measure of bark beetle resistance, depended on season of measurement. Trees in both thinning treatments continued to have increased leaf toughness, a measure of resistance to insect folivores. Our results show that many beneficial effects of restoration treatments on carbon, water, and nitrogen relations and insect‐resistance characteristics of pre‐settlement ponderosa pines continue to be expressed 7 years after treatment at the GPNA.     

Radial and stand‐level thinning treatments: 15‐year growth response of legacy ponderosa and Jeffrey pine trees

Restoration efforts to improve vigor of large, old trees and decrease risk to high‐intensity wildland fire and drought‐mediated insect mortality often include reductions in stand density. We examined 15‐year growth response of old ponderosa pine (Pinus ponderosa) and Jeffrey pine (Pinus jeffreyi) trees in northeastern California, U.S.A. to two levels of thinning treatments compared to an untreated (control) area. Density reductions involved radial thinning (thinning 9.1 m around individual trees) and stand thinning. Annual tree growth in the stand thinning increased immediately following treatment and was sustained over the 15 years. In contrast, radial thinning did not increase growth, but slowed decline compared to control trees. Available soil moisture was higher in the stand thinning than the control for 5 years post‐treatment and likely extended seasonal tree growth. Our results show that large, old trees can respond to restoration thinning treatments, but that the level of thinning impacts this response. Stand thinning must be sufficiently intensive to improve old tree growth and health, in part due to increasing available soil moisture. Importantly, focusing stand density reductions around the immediate neighborhood of legacy trees was insufficient to elicit a growth response, calling into question treatments attempting to increase vigor of legacy trees while still maintaining closed canopies in dry, coniferous forest types. Although radial thinning did not affect tree growth rates, this treatment may still achieve other resource objectives not studied here, such as protecting wildlife habitat, reducing the risk of severe fire injury, and decreasing susceptibility to bark beetle attacks.     

Construction cost of loblolly and ponderosa pine leaves grown with varying carbon and nitrogen availability

We grew loblolly and ponderosa pine seedlings in a factorial experiment with two CO₂ partial pressures (35 and 70 Pa), and two nitrogen treatments (1.0 and 3.5 mol m^?3 NH₄⁺), for one growing season to examine the effects of carbon and nitrogen availability on leaf construction cost. Growth in elevated CO₂ reduced leaf nitrogen concentrations by 17 to 40%, and increased C:N by 22 to 68%. Elevated N availability increased leaf N concentrations and decreased C:N. Non-structural carbohydrates increased in high-CO₂-grown loblolly seedlings, except in fascicles from low N, and in ponderosa primary and fascicle leaves grown in high N. In loblolly, increases in starch were nearly 2-fold greater than the increases in soluble sugars. In ponderosa, only the soluble sugars were affected by CO₂. Leaf construction cost (g glucose g^?1 dm) varied by 9.3% across all treatments. All of the variation in loblolly leaf construction cost could be explained by changes in non-structural carbohydrates. A model of the response of construction cost to changes in the mass of different biochemical fractions suggests that the remainder of the variation in ponderosa, not explained by non-structural carbohydrates, is probably attributable to changes in lignin, phenolic or protein concentrations.     

Common and Uncommon Understory Species Differentially Respond to Restoration Treatments in Ponderosa Pine/Douglas-Fir Forests, Montana

Restoration treatments have been widely advocated to address declining conditions in Pinus ponderosa forests throughout the western United States. However, few studies have examined treatment effects on individual plant species or whether responses differ for common species and uncommon species (those with low abundance in the community)—information that may be critical in managing for long‐term biodiversity. We investigated understory species responses to restoration treatments in ponderosa pine/Douglas‐fir forests using a randomized block experimental design with three blocks and four treatments (control, burn‐only, thin‐only, and thin‐burn). Understory vegetation was sampled before treatment and for three consecutive years after treatment. We used richness and an index of uniqueness to compare responses of common and uncommon native understory species among treatments, and indicator species analysis to identify individual species that responded to each treatment. Treatments that included thinning had significantly more unique species assemblages than the control. The thin‐only treatment increased common native species richness, whereas all active treatments significantly increased uncommon native species richness over the control, especially the thin‐burn. Generally, life‐forms did not explain the responses of individual species, though in the final sampling year several graminoids were exclusively indicative of treatments that included thinning. Very few species had reduced abundance in the thinning and burning treatments by the final sample year, whereas many uncommon and short‐lived species benefited from active treatments, especially the combined thin‐burn treatment. Active restoration treatments in these forests may foster plant diversity by minimally impacting common species while significantly benefiting disturbance‐dependent native species.     

Ethanol synthesis, nitrogen, carbohydrates, and growth in tissues from nitrogen fertilized Pseudotsuga menziesii (Mirb.) Franco and Pinus ponderosa Dougl. ex Laws. seedlings

 Seedlings of Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, and ponderosa pine, Pinus ponderosa Dougl. ex Laws., were grown in a controlled environment and fertilized with nutrient solutions containing 150 ppm (+N), or 0 ppm nitrogen (−N). These treatments greatly altered seedling growth, and the concentrations of N and carbohydrates in their tissues. Metabolically active tissues, such as roots, incubated with a limited supply of O₂ became hypoxic faster and synthesized more ethanol than less active tissues, such as needles. All tissues that were incubated for 4 h in N₂ synthesized ethanol. Needles incubated in N₂ and light had much lower quantities of ethanol than needles in N₂ and dark, suggesting that O₂ from photosynthetsis limited internal anoxia. Most tissues from +N seedlings synthesized greater quantities of ethanol in N₂ anoxia than tissues from −N seedlings, probably because they were able to produce more enzymes with a greater availability of N. However, this increase in ethanol synthesis between N treatments was most pronounced in the phloem. Ethanol and soluble sugar concentrations were negatively related in needles and positively related in roots of N+ seedlings, but not −N seedlings. Starch concentrations had no effect on the amount of ethanol produced by any tissue. Regardless of N treatments, all tissues from ponderosa pine produced more N₂-induced ethanol than Douglas-fir, in part because its tissues contained different concentrations of soluble sugars and N as a consequence of phenological differences between the species. However, ponderosa pine tissues may also maintain greater quantities of anaerobic enzymes, or their isozymes than Douglas-fir. Received: 22 February 1998 / Accepted 23 June 1998     

Effects of Climate Variability and Accelerated Forest Thinning on Watershed-Scale Runoff in Southwestern USA Ponderosa Pine Forests

The recent mortality of up to 20% of forests and woodlands in the southwestern United States, along with declining stream flows and projected future water shortages, heightens the need to understand how management practices can enhance forest resilience and functioning under unprecedented scales of drought and wildfire. To address this challenge, a combination of mechanical thinning and fire treatments are planned for 238,000 hectares (588,000 acres) of ponderosa pine (Pinus ponderosa) forests across central Arizona, USA. Mechanical thinning can increase runoff at fine scales, as well as reduce fire risk and tree water stress during drought, but the effects of this practice have not been studied at scales commensurate with recent forest disturbances or under a highly variable climate. Modifying a historical runoff model, we constructed scenarios to estimate increases in runoff from thinning ponderosa pine at the landscape and watershed scales based on driving variables: pace, extent and intensity of forest treatments and variability in winter precipitation. We found that runoff on thinned forests was about 20% greater than unthinned forests, regardless of whether treatments occurred in a drought or pluvial period. The magnitude of this increase is similar to observed declines in snowpack for the region, suggesting that accelerated thinning may lessen runoff losses due to warming effects. Gains in runoff were temporary (six years after treatment) and modest when compared to mean annual runoff from the study watersheds (0–3%). Nonetheless gains observed during drought periods could play a role in augmenting river flows on a seasonal basis, improving conditions for water-dependent natural resources, as well as benefit water supplies for downstream communities. Results of this study and others suggest that accelerated forest thinning at large scales could improve the water balance and resilience of forests and sustain the ecosystem services they provide.     

Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

Effects of CO2 and nitrogen fertilization on vegetation and soil nutrient content in juvenile ponderosa pine

This paper summarizes the data on nutrient uptake and soil responses in opentop chambers planted with ponderosa pine (Pinus ponderosa Laws.) treated with both N and CO₂. Based upon the literature, we hypothesized that 1) elevated CO₂ would cause increased growth and yield of biomass per unit uptake of N even if N is limiting, and 2) elevated CO₂ would cause increased biomass yield per unit uptake of other nutrients only by growth dilution and only if they are non-limiting. Hypothesis 1 was supported only in part: there were greater yields of biomass per unit N uptake in the first two years of growth but not in the third year. Hypothesis 2 was supported in many cases: elevated CO₂ caused growth dilution (decreased concentrations but not decreased uptake) of P, S, and Mg. Effects of elevated CO₂ on K, Ca, and B concentrations were smaller and mostly non-significant. There was no evidence that N responded in a unique manner to elevated CO₂, despite its unique role in rubisco. Simple growth dilution seemed to explain nutrient responses in almost all cases.There were significant declines in soil exchangeable K⁺, Ca²⁺, Mg²⁺ and extractable P over time which were attributed to disturbance effects associated with plowing. The only statistically significant treatment effects on soils were negative effects of elevated CO₂ on mineralizeable N and extractable P, and positive effects of both N fertilization and CO₂ on exchangeable Al³⁺. Soil exchangeable K⁺, Ca²⁺, and Mg²⁺ pools remained much higher than vegetation pools, but extractable P pools were lower than vegetation pools in the third year of growth. There were also large losses of both native soil N and fertilizer N over time. These soil N losses could account for the observed losses in exchangeable K⁺, Ca²⁺, Mg²⁺ if N was nitrified and leached as NO ₃ ^– .     

Wildfire effects on carbon and nitrogen in inland coniferous forests

A ponderosa pine/Douglas-fir forest (Pinus ponderosa Dougl., Pseudotsuga menziesii (Mirb.) Franco; PP/DF) and a lodgepole pine/Engelmann spruce forest (Pinus contorta Loud., Picea engelmannii Parry ex Engelm.; LP/ES) located on the eastern slopes of the Cascade Mountains in Washington state, USA, were examined following severe wildfire to compare total soil carbon and nitrogen capitals with unburned (control) forests. One year after fire, the average C content (60 cm depth) of PP/DF and LP/ES soil was 30% (25 Mg ha^-1) and 10% (7 Mg ha^-1) lower than control soil. Average N content on the burned PP/DF and LP/ES plots was 46% (3.0 Mg ha^-1) and 13% (0.4 Mg ha^-1) lower than control soil. The reduction in C and N in the PP/DF soil was largely the result of lower nutrient capitals in the burned Bw horizons (12–60 cm depth) relative to control plots. It is unlikely that the 1994 fire substantially affected nutrient capitals in the Bw horizons; however, natural variability or past fire history could be responsible for the varied nutrient capitals observed in the subsurface soils. Surface erosion (sheet plus rill) removed between 15 and 18 Mg ha^-1 of soil from the burned plots. Nutrient losses through surface erosion were 280 kg C ha^-1 and 14 kg N ha^-1 in the PP/DF, whereas LP/ES losses were 640 and 22 kg ha^-1 for C and N, respectively. In both forests, surface erosion of C and N was _∼1% to 2% of the A-horizon capital of these elements in unburned soil. A bioassay (with lettuce as an indicator plant) was used to compare soils from low-, moderate- and high-severity burn areas relative to control soil. In both forests, low-severity fire increased lettuce yield by 70–100% of controls. With more severe fire, yield decreased in the LP/ES relative to the low-intensity burn soil; however, only in the high-severity treatment was yield reduced (14%) from the control. Moderate- and high-severity burn areas in the PP/DF were fertilized with _∼56 kg ha^-1 of N four months prior to soil sampling. In these soils, yield was 70–80% greater than the control. These results suggest that short-term site productivity can be stimulated by low-severity fire, but unaffected or reduced by more severe fire in the types of forests studied. Post-fire fertilization with N could increase soil productivity where other environmental factors do not limit growth. This revised version was published online in June 2006 with corrections to the Cover Date.     

Soil N mineralization and nitrification in relation to nitrogen solution chemistry in a small forested watershed

Spatial variations in soil processes regulating mineral N losses to streams were studied in a small watershed near Toronto, Ontario. Annual net N mineralization in the 0–8 cm soil was measured in adjacent upland and riparian forest stands using in situ soil incubations from April 1985 to 1987. Mean annual rates of soil N mineralization and nitrification were higher in a maple soil (93.8 and 87.0 kg.ha^–1) than in a pine soil (23.3 and 8.2 kg.ha^–1 ). Very low mean rates of mineralization (3.3 kg.ha^–1) and nitrification (3.4 kg.ha^–1) were found in a riparian hemlock stand. Average NO₃-N concentrations in soil solutions were 0.3–1.0 mg.L^–1 in the maple stand and >0.06mg.L^–1 in the pine stand. Concentrations of NO₃–N in shallow ground water and stream water were 3–4× greater in a maple subwatershed than in a pine subwatershed. Rapid N uptake by vegetation was an important mechanism reducing solution losses of NO₃–N in the maple stand. Low rates of nitrification were mainly responsible for negligible NO₃–N solution losses in the pine stand.