Feast not famine: Nitrogen pools recover rapidly in 25-yr-old postfire lodgepole pine.

The extent of young postfire conifer forests is growing throughout western North America as the frequency and size of high-severity fires increase, making it important to understand ecosystem structure and function in early seral forests. Understanding nitrogen (N) dynamics during postfire stand development is especially important because northern conifers are often N limited. We resampled lodgepole pine (Pinus contorta var. latifolia) stands that regenerated naturally after the 1988 fires in Yellowstone National Park (Wyoming, USA) to ask (1) How have N pools and fluxes changed over a decade (15 to 25 yr postfire) of very rapid forest growth? (2) At postfire year 25, how do N pools and fluxes vary with lodgepole pine density and productivity? Lodgepole pine foliage, litter (annual litterfall, forest-floor litter), and mineral soils were sampled in 14 plots (0.25 ha) that varied in postfire lodgepole pine density (1,500 to 344,000 stems/ha) and aboveground net primary production (ANPP; 1.4 to 16.1 Mg·ha-1 ·yr-1 ). Counter to expectation, foliar N concentrations in lodgepole pine current-year and composite needles (1.33 and 1.11% N, respectively) had not changed over time. Further, all measured ecosystem N pools increased substantially: foliar N increased to 89 kg N/ha (+93%), O-horizon N increased to 39 kg N/ha (+38%), and mineral soil percent total N (0-15 cm) increased to 0.08% (+33%). Inorganic N availability also increased to 0.69 µg N·[g resin]-1 ·d-1 (+165%). Thus, soil N did not decline as live biomass N pools increased. Among stands, biomass N pools at postfire year 25 remained strongly influenced by early postfire tree density: foliar and litterfall N concentrations declined with lodgepole pine density and ANPP, but the foliar N pool increased. Lodgepole pine ANPP correlated negatively with annual resin-sorbed N, and we found no indication of widespread N limitation. The large increases in N pools cannot be explained by atmospheric N deposition or presence of known N fixers. These results suggest an unmeasured N source and are consistent with recent reports of N fixation in young lodgepole pine.

Twenty-four years after theYellowstone Fires: Are postfire lodgepole pine stands converging in structure and function?

Disturbance and succession have long been of interest in ecology, but how landscape patterns of ecosystem structure and function evolve following large disturbances is poorly understood. After nearly 25 years, lodgepole pine (Pinus contorta var. latifolia) forests that regenerated after the 1988 Yellowstone Fires (Wyoming, USA) offer a prime opportunity to track the fate of disturbance-created heterogeneity in stand structure and function in a wilderness setting. In 2012, we resampled 72 permanent plots to ask (1) How have postfire stand structure and function changed between 11 and 24 yr postfire, and what variables explain these patterns and changes? (2) How has landscape-level (among-stand) variability in postfire stand structure and function changed between 11 and 24 yr postfire? We expected to see evidence of convergence beginning to emerge, but also that initial postfire stem density would still determine trajectories of biomass accumulation. After 24 yr, postfire lodgepole pine density remained very high (mean = 21,738 stems/ha, range = 0-344,067 stems/ha). Stem density increased in most plots between 11 and 24 yr postfire, but declined sharply where 11-yr-postfire stem density was > 72,000 stems/ha. Stems were small in high-density stands, but stand-level lodgepole pine leaf area, foliage biomass, and live aboveground biomass increased over time and with increasing stem density. After 24 yr, mean annual lodgepole pine aboveground net primary production (ANPP) was high (mean = 5 Mg · ha⁻¹ · yr⁻¹, range = 0-16.5 Mg · ha⁻¹ · yr⁻¹). Among stands, lodgepole pine ANPP increased with stem density, which explained 69% of the variation; another 8% of the variation was explained by environmental covariates. Early patterns of postfire lodgepole pine regeneration, which were contingent on prefire serotiny and fire severity, remained the dominant driver of stand structure and function. We observed mechanisms that would lead to convergence in stem density (structure) over time, but it was landscape variation in functional variables that declined substantially. Stand structure and function have not converged across the burned landscape, but our evidence suggests function will converge sooner than structure.

Landscape variation in tree regeneration and snag fall drive fuel loads in 24-year old post-fire lodgepole pine forests.

Escalating wildfire in subalpine forests with stand-replacing fire regimes is increasing the extent of early-seral forests throughout the western USA. Post-fire succession generates the fuel for future fires, but little is known about fuel loads and their variability in young post-fire stands. We sampled fuel profiles in 24-year-old post-fire lodgepole pine (Pinus contorta var. latifolia) stands (n = 82) that regenerated from the 1988 Yellowstone Fires to answer three questions. (1) How do canopy and surface fuel loads vary within and among young lodgepole pine stands? (2) How do canopy and surface fuels vary with pre- and post-fire lodgepole pine stand structure and environmental conditions? (3) How have surface fuels changed between eight and 24 years post-fire? Fuel complexes varied tremendously across the landscape despite having regenerated from the same fires. Available canopy fuel loads and canopy bulk density averaged 8.5 Mg/ha (range 0.0-46.6) and 0.24 kg/m3 (range: 0.0-2.3), respectively, meeting or exceeding levels in mature lodgepole pine forests. Total surface-fuel loads averaged 123 Mg/ha (range: 43-207), and 88% was in the 1,000-h fuel class. Litter, 1-h, and 10-h surface fuel loads were lower than reported for mature lodgepole pine forests, and 1,000-h fuel loads were similar or greater. Among-plot variation was greater in canopy fuels than surface fuels, and within-plot variation was greater than among-plot variation for nearly all fuels. Post-fire lodgepole pine density was the strongest positive predictor of canopy and fine surface fuel loads. Pre-fire successional stage was the best predictor of 100-h and 1,000-h fuel loads in the post-fire stands and strongly influenced the size and proportion of sound logs (greater when late successional stands had burned) and rotten logs (greater when early successional stands had burned). Our data suggest that 76% of the young post-fire lodgepole pine forests have 1,000-h fuel loads that exceed levels associated with high-severity surface fire potential, and 63% exceed levels associated with active crown fire potential. Fire rotations in Yellowstone National Park are predicted to shorten to a few decades and this prediction cannot be ruled out by a lack of fuels to carry repeated fires.

Tree recruitment in relation to climate and fire in northern Mexico.

Extensive changes in montane forest structure have occurred throughout the U.S. Southwest following Euro-American settlement. These changes are a product of confounding effects of disturbance, climate variability, species competition, and modern land use changes. Pronounced forest reproduction events in the Southwest have generally occurred in climatically wet periods but have also followed widespread fire exclusion. Understanding the ecological processes driving such events has important implications for forest restoration, although these efforts remain difficult due to confounding factors. Separation of these interacting factors was possible in the Sierra San Luis of northern Mexico where we investigated climate, fire, and tree recruitment in areas with continued frequent fires or where fire exclusion came relatively late (1940s). Fires were strongly tied to interannual wet-dry cycles of climate, whereas recruitment peaks were more closely tied to local processes, namely, fire-free periods, than to broad-scale climatically wet conditions. The greatest pulse of tree recruitment coincided with a pronounced mid-century drought (1942-1957) and a period of reduced fire frequency. The second largest pulse of recruitment (ca. 1900) preceded a well-documented period of recruitment (and an anomalously wet period) elsewhere across the Southwest in the 1910s-1920s, and also coincided with specific fire-free periods during below-average precipitation. We also found greater spatial dependence and clustering in older age classes of trees. This spatial pattern indicates a legacy of fire-induced mortality in shaping stand structure, underscoring the importance of frequent fire effects on spatial variability in forests.

Fire severity and tree regeneration following bark beetle outbreaks: the role of outbreak stage and burning conditions.

The degree to which recent bark beetle (Dendroctonus ponderosae) outbreaks may influence fire severity and postfire tree regeneration is of heightened interest to resource managers throughout western North America, but empirical data on actual fire effects are lacking. Outcomes may depend on burning conditions (i.e., weather during fire), outbreak severity, or intervals between outbreaks and subsequent fire. We studied recent fires that burned through green-attack/red-stage (outbreaks <3 years before fire) and gray-stage (outbreaks 3­15 years before fire) subalpine forests dominated by lodgepole pine (Pinus contorta var. latifolia) in Greater Yellowstone, Wyoming, USA, to determine if fire severity was linked to prefire beetle outbreak severity and whether these two disturbances produced compound ecological effects on postfire tree regeneration. With field data from 143 postfire plots that burned under different conditions, we assessed canopy and surface fire severity, and postfire tree seedling density against prefire outbreak severity. In the green-attack/red stage, several canopy fire-severity measures increased with prefire outbreak severity under moderate burning conditions. Under extreme conditions, few fire-severity measures were related to prefire outbreak severity, and effect sizes were of marginal biological significance. The percentage of tree stems and basal area killed by fire increased with more green-attack vs. red-stage trees (i.e., the earliest stages of outbreak). In the gray stage, by contrast, most fire-severity measures declined with increasing outbreak severity under moderate conditions, and fire severity was unrelated to outbreak severity under extreme burning conditions. Postfire lodgepole pine seedling regeneration was unrelated to prefire outbreak severity in either post-outbreak stage, but increased with prefire serotiny. Results suggest bark beetle outbreaks can affect fire severity in subalpine forests under moderate burning conditions, but have little effect on fire severity under extreme burning conditions when most large wildfires occur in this system. Thus, beetle outbreak severity was moderately linked to fire severity, but the strength and direction of the linkage depended on both endogenous (outbreak stage) and exogenous (fire weather) factors. Closely timed beetle outbreak and fire did not impart compound effects on tree regeneration, suggesting the presence of a canopy seedbank may enhance resilience to their combined effects.

Continued warming could transform Greater Yellowstone fire regimes by mid-21st century.

Climate change is likely to alter wildfire regimes, but the magnitude and timing of potential climate-driven changes in regional fire regimes are not well understood. We considered how the occurrence, size, and spatial location of large fires might respond to climate projections in the Greater Yellowstone ecosystem (GYE) (Wyoming), a large wildland ecosystem dominated by conifer forests and characterized by infrequent, high-severity fire. We developed a suite of statistical models that related monthly climate data (1972-1999) to the occurrence and size of fires >200 ha in the northern Rocky Mountains; these models were cross-validated and then used with downscaled (~12 km × 12 km) climate projections from three global climate models to predict fire occurrence and area burned in the GYE through 2099. All models predicted substantial increases in fire by midcentury, with fire rotation (the time to burn an area equal to the landscape area) reduced to <30 y from the historical 100-300 y for most of the GYE. Years without large fires were common historically but are expected to become rare as annual area burned and the frequency of regionally synchronous fires increase. Our findings suggest a shift to novel fire-climate-vegetation relationships in Greater Yellowstone by midcentury because fire frequency and extent would be inconsistent with persistence of the current suite of conifer species. The predicted new fire regime would transform the flora, fauna, and ecosystem processes in this landscape and may indicate similar changes for other subalpine forests.

Influence of recent bark beetle outbreak on fire severity and postfire tree regeneration in montane Douglas-fir forests.

Understanding how disturbances interact to shape ecosystems is a key challenge in ecology. In forests of western North America, the degree to which recent bark beetle outbreaks and subsequent fires may be linked (e.g., outbreak severity affects fire severity) and/ or whether these two disturbances produce compound effects on postfire succession is of widespread interest. These interactions remain unresolved, largely because field data from actual wildfires following beetle outbreaks are lacking. We studied the 2008 Gunbarrel Fire, which burned 27 200 ha in Douglas-fir (Pseudotsuga menziesii) forests that experienced a bark beetle outbreak 4-13 years prefire ("gray stage," after trees have died and needles have dropped), to determine whether outbreak severity influenced subsequent fire severity and postfire tree regeneration. In 85 sample plots we recorded prefire stand structure and outbreak severity; multiple measures of canopy and forest-floor fire severity; and postfire tree seedling density. Prefire outbreak severity was not related to any measure of fire severity except for mean bole scorch, which declined slightly with increasing outbreak severity. Instead, fire severity varied with topography and burning conditions (proxy for weather at time of fire). Postfire Douglas-fir regeneration was low, with tree seedlings absent in 65% of plots. Tree seedlings were abundant in plots of low fire severity that also had experienced low outbreak severity (mean = 1690 seedlings/ha), suggesting a dual filter on tree regeneration. Although bark beetles and fire collectively reduced live basal area to < 5% and increased snag density to > 2000% of pre-outbreak levels, the lack of relationship between beetle outbreak and fire severity suggests that these disturbances were not linked. Nonetheless, effects on postfire tree regeneration suggest compound disturbance interactions that contribute to the structural heterogeneity characteristic of mid/lower montane forests.

Bark beetle effects on fuel profiles across a range of stand structures in Douglas-fir forests of Greater Yellowstone.

Consequences of bark beetle outbreaks for forest wildfire potential are receiving heightened attention, but little research has considered ecosystems with mixed-severity fire regimes. Such forests are widespread, variable in stand structure, and often fuel limited, suggesting that beetle outbreaks could substantially alter fire potentials. We studied canopy and surface fuels in interior Douglas-fir (Pseudotsuga menziesii v. glauca) forests in Greater Yellowstone, Wyoming, USA, to determine how fuel characteristics varied with time since outbreak of the Douglas-fir beetle (Dendroctonus pseudotsugae). We sampled five stands in each of four outbreak stages, validated for pre-outbreak similarity: green (undisturbed), red (1-3 yr), gray (4-14 yr), and silver (25-30 yr). General linear models were used to compare variation in fuel profiles associated with outbreak to variation associated with the range of stand structures (dense mesic forest to open xeric parkland) characteristic of interior Douglas-fir forest. Beetle outbreak killed 38-83% of basal area within stands, generating a mix of live trees and snags over several years. Canopy fuel load and bulk density began declining in the red stage via needle drop and decreased by approximately 50% by the silver stage. The dead portion of available canopy fuels peaked in the red stage at 41%. After accounting for background variation, there was little effect of beetle outbreak on surface fuels, with differences mainly in herbaceous biomass (50% greater in red stands) and coarse woody fuels (doubled in silver stands). Within-stand spatial heterogeneity of fuels increased with time since outbreak, and surface-to-crown continuity decreased and remained low because of slow/sparse regeneration. Collectively, results suggest reduced fire potentials in post-outbreak stands, particularly for crown fire after the red stage, although abundant coarse fuels in silver stands may increase burn residence time and heat release. Outbreak effects on fuels were comparable to background variation in stand structure. The net effect of beetle outbreak was to shift the structure of mnsic closed-canopy stands toward that of parklands, and to shift xeric parklands toward very sparse woodlands. This study highlights the importance of evaluating outbreak effects in the context of the wide structural variation inherent to many forest types in the absence of beetle disturbance.

Reconciling divergent interpretations of quaking aspen decline on the northern Colorado Front Range.

Ecologists have debated over the past 65 years whether quaking aspen (Populus tremuloides Michx.) has or has not declined in abundance, vigor, or regeneration in western North America. Many studies have provided divergent interpretations of the condition of aspen forests, leading to difficulty in translating this ecological information into management recommendations. To reconcile these contrasting conclusions and to test the hypothesis that multiple types of aspen decline and persistence occur simultaneously on heterogeneous landscapes, we assessed 91 aspen stands across the northern Colorado Front Range to determine the range of ecological conditions that underlie aspen decline or persistence. Approximately 15% of aspen forest area in our sample exhibited dieback of mature stems coupled with a lack of young trees indicative of declining stands, most often at lower elevations where elk browsing is heavy and chronic, and where effects of fire exclusion have been most significant. However, 52% of the area sampled had multiple cohorts indicative of self-replacing or persistent stands. Conifer dominance was increasing in over 33% of all aspen forest area sampled, most often at high elevations among lodgepole pine (Pinus contorta var. latifolia Englem. ex Wats.) forests. Reconstructions of relative basal area and density of aspen and lodgepole pine in these stands suggest cyclical dominance of these species, where conifers gradually replace aspen over long fire intervals, and aspen vigorously re-establish following stand-replacing fires. The diversity of ecological contexts across the northern Colorado Front Range creates a variety of aspen dynamics leading to decline or persistence, and no single trend describes the general condition of aspen forests in appropriate detail for managers. Active management may be useful in preserving individual stands at fine scales, but management prescriptions should reflect specific drivers of decline in these stands.

Post-fire spatial patterns of soil nitrogen mineralization and microbial abundance.

Stand-replacing fires influence soil nitrogen availability and microbial community composition, which may in turn mediate post-fire successional dynamics and nutrient cycling. However, fires create patchiness at both local and landscape scales and do not result in consistent patterns of ecological dynamics. The objectives of this study were to (1) quantify the spatial structure of microbial communities in forest stands recently affected by stand-replacing fire and (2) determine whether microbial variables aid predictions of in situ net nitrogen mineralization rates in recently burned stands. The study was conducted in lodgepole pine (Pinus contorta var. latifolia) and Engelmann spruce/subalpine fir (Picea engelmannii/Abies lasiocarpa) forest stands that burned during summer 2000 in Greater Yellowstone (Wyoming, USA). Using a fully probabilistic spatial process model and Bayesian kriging, the spatial structure of microbial lipid abundance and fungi-to-bacteria ratios were found to be spatially structured within plots two years following fire (for most plots, autocorrelation range varied from 1.5 to 10.5 m). Congruence of spatial patterns among microbial variables, in situ net N mineralization, and cover variables was evident. Stepwise regression resulted in significant models of in situ net N mineralization and included variables describing fungal and bacterial abundance, although explained variance was low (R²<0.29). Unraveling complex spatial patterns of nutrient cycling and the biotic factors that regulate it remains challenging but is critical for explaining post-fire ecosystem function, especially in Greater Yellowstone, which is projected to experience increased fire frequencies by mid 21(st) Century.

Inorganic nitrogen availability after severe stand-replacing fire in the Greater Yellowstone ecosystem.

Understanding ecosystem processes as they relate to wildfire and vegetation dynamics is of growing importance as fire frequency and extent increase throughout the western United States. However, the effects of severe, stand-replacing wildfires are poorly understood. We studied inorganic nitrogen pools and mineralization rates after stand-replacing wildfires in the Greater Yellowstone Ecosystem, Wyoming. After fires that burned in summer 2000, soil ammonium concentration peaked in 2001 (33 mg NH(4)-N x kg(soil)(-1)); soil nitrate increased subsequently (2.7 mg NO(3)-N.kg(soil)(-1) in 2003) but was still low. However, annual net ammonification rates were largely negative from 2001 to 2004, indicating ammonium depletion. Thus, although net nitrification rates were positive, annual net nitrogen mineralization (net ammonification plus net nitrification) remained low. Aboveground net primary production (ANPP) increased from 0.25 to 1.6 Mg x ha(-1) x yr(-1) from 2001 to 2004, but variation in ANPP among stands was not related to net nitrogen mineralization rates. Across a broader temporal gradient (stand age zero to >250 yr), negative rates of net annual ammonification were especially pronounced in the first postfire year. Laboratory incubations using (15)N isotope pool dilution revealed that gross production of ammonium was reduced and ammonium consumption greatly exceeded gross production during the initial postfire years. Our results suggest a microbial nitrogen sink for several years after severe, stand-replacing fire, confirming earlier hypotheses about postdisturbance succession and nutrient cycling in cold, fire-dominated coniferous forests. Postfire forests in Yellowstone seem to be highly conservative for nitrogen, and microbial immobilization of ammonium plays a key role during early succession.

Regional vegetation die-off in response to global-change-type drought.

Future drought is projected to occur under warmer temperature conditions as climate change progresses, referred to here as global-change-type drought, yet quantitative assessments of the triggers and potential extent of drought-induced vegetation die-off remain pivotal uncertainties in assessing climate-change impacts. Of particular concern is regional-scale mortality of overstory trees, which rapidly alters ecosystem type, associated ecosystem properties, and land surface conditions for decades. Here, we quantify regional-scale vegetation die-off across southwestern North American woodlands in 2002-2003 in response to drought and associated bark beetle infestations. At an intensively studied site within the region, we quantified that after 15 months of depleted soil water content, >90% of the dominant, overstory tree species (Pinus edulis, a piñon) died. The die-off was reflected in changes in a remotely sensed index of vegetation greenness (Normalized Difference Vegetation Index), not only at the intensively studied site but also across the region, extending over 12,000 km2 or more; aerial and field surveys confirmed the general extent of the die-off. Notably, the recent drought was warmer than the previous subcontinental drought of the 1950s. The limited, available observations suggest that die-off from the recent drought was more extensive than that from the previous drought, extending into wetter sites within the tree species' distribution. Our results quantify a trigger leading to rapid, drought-induced die-off of overstory woody plants at subcontinental scale and highlight the potential for such die-off to be more severe and extensive for future global-change-type drought under warmer conditions.