Toward Meaningful Snag-Management Guidelines for Postfire Salvage Logging in North American Conifer Forests

Abstract:  The bird species in western North America that are most restricted to, and therefore most dependent on, severely burned conifer forests during the first years following a fire event depend heavily on the abundant standing snags for perch sites, nest sites, and food resources. Thus, it is critical to develop and apply appropriate snag-management guidelines to implement postfire timber harvest operations in the same locations. Unfortunately, existing guidelines designed for green-tree forests cannot be applied to postfire salvage sales because the snag needs of snag-dependent species in burned forests are not at all similar to the snag needs of snag-dependent species in green-tree forests. Birds in burned forests have very different snag-retention needs from those cavity-nesting bird species that have served as the focus for the development of existing snag-management guidelines. Specifically, many postfire specialists use standing dead trees not only for nesting purposes but for feeding purposes as well. Woodpeckers, in particular, specialize on wood-boring beetle larvae that are superabundant in fire-killed trees for several years following severe fire. Species such as the Black-backed Woodpecker ( Picoides arcticus ) are nearly restricted in their habitat distribution to severely burned forests. Moreover, existing postfire salvage-logging studies reveal that most postfire specialist species are completely absent from burned forests that have been (even partially) salvage logged. I call for the long-overdue development and use of more meaningful snag-retention guidelines for postfire specialists, and I note that the biology of the most fire-dependent bird species suggests that even a cursory attempt to meet their snag needs would preclude postfire salvage logging in those severely burned conifer forests wherein the maintenance of biological diversity is deemed important.     

Potential shifts in dominant forest cover in interior Alaska driven by variations in fire severity

Large fire years in which >1% of the landscape burns are becoming more frequent in the Alaskan (USA) interior, with four large fire years in the past 10 years, and 79 000 km2 (17% of the region) burned since 2000. We modeled fire severity conditions for the entire area burned in large fires during a large fire year (2004) to determine the factors that are most important in estimating severity and to identify areas affected by deep-burning fires. In addition to standard methods of assessing severity using spectral information, we incorporated information regarding topography, spatial pattern of burning, and instantaneous characteristics such as fire weather and fire radiative power. Ensemble techniques using regression trees as a base learner were able to determine fire severity successfully using spectral data in concert with other relevant geospatial data. This method was successful in estimating average conditions, but it underestimated the range of severity. This new approach was used to identify black spruce stands that experienced intermediate- to high-severity fires in 2004 and are therefore susceptible to a shift in regrowth toward deciduous dominance or mixed dominance. Based on the output of the severity model, we estimate that 39% (approximately 4000 km2) of all burned black spruce stands in 2004 had <10 cm of residual organic layer and may be susceptible a postfire shift in plant functional type dominance, as well as permafrost loss. If the fraction of area susceptible to deciduous regeneration is constant for large fire years, the effect of such years in the most recent decade has been to reduce black spruce stands by 4.2% and to increase areas dominated or co-dominated by deciduous forest stands by 20%. Such disturbance-driven modifications have the potential to affect the carbon cycle and climate system at regional to global scales.     

Fire severity and ecosytem responses following crown fires in California shrublands

Chaparral shrublands burn in large high-intensity crown fires. Managers interested in how these wildfires affect ecosystem processes generally rely on surrogate measures of fire intensity known as fire severity metrics. In shrublands burned in the autumn of 2003, a study of 250 sites investigated factors determining fire severity and ecosystem responses. Using structural equation modeling we show that stand age, prefire shrub density, and the shortest interval of the prior fire history had significant direct effects on fire severity, explaining > 50% of the variation in severity. Fire severity per se is of interest to resource managers primarily because it is presumed to be an indicator of important ecosystem processes such as vegetative regeneration, community recovery, and erosion. Fire severity contributed relatively little to explaining patterns of regeneration after fire. Two generalizations can be drawn: fire severity effects are mostly shortlived, i.e., by the second year they are greatly diminished, and fire severity may have opposite effects on different functional types. Species richness exhibited a negative relationship to fire severity in the first year, but fire severity impacts were substantially less in the second postfire year and varied by functional type. Much of this relationship was due to alien plants that are sensitive to high fire severity; at all scales from 1 to 1000 m2, the percentage of alien species in the postfire flora declined with increased fire severity. Other aspects of disturbance history are also important determinants of alien cover and richness as both increased with the number of times the site had burned and decreased with time since last fire. A substantial number of studies have shown that remote-sensing indices are correlated with field measurements of fire severity. Across our sites, absolute differenced normalized burn ratio (dNBR) was strongly correlated with field measures of fire severity and with fire history at a site but relative dNBR was not. Despite being correlated with fire severity, absolute dNBR showed little or no relationship with important ecosystem responses to wildfire such as shrub resprouting or total vegetative regeneration. These findings point to a critical need for further research on interpreting remote sensing indices as applied to postfire management of these shrublands.     

Rapid Increases and Time‐Lagged Declines in Amphibian Occupancy after Wildfire

Climate change is expected to increase the frequency and severity of drought and wildfire. Aquatic and moisture‐sensitive species, such as amphibians, may be particularly vulnerable to these modified disturbance regimes because large wildfires often occur during extended droughts and thus may compound environmental threats. However, understanding of the effects of wildfires on amphibians in forests with long fire‐return intervals is limited. Numerous stand‐replacing wildfires have occurred since 1988 in Glacier National Park (Montana, U.S.A.), where we have conducted long‐term monitoring of amphibians. We measured responses of 3 amphibian species to fires of different sizes, severity, and age in a small geographic area with uniform management. We used data from wetlands associated with 6 wildfires that burned between 1988 and 2003 to evaluate whether burn extent and severity and interactions between wildfire and wetland isolation affected the distribution of breeding populations. We measured responses with models that accounted for imperfect detection to estimate occupancy during prefire (0–4 years) and different postfire recovery periods. For the long‐toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), occupancy was not affected for 6 years after wildfire. But 7–21 years after wildfire, occupancy for both species decreased ≥25% in areas where >50% of the forest within 500 m of wetlands burned. In contrast, occupancy of the boreal toad (Anaxyrus boreas) tripled in the 3 years after low‐elevation forests burned. This increase in occupancy was followed by a gradual decline. Our results show that accounting for magnitude of change and time lags is critical to understanding population dynamics of amphibians after large disturbances. Our results also inform understanding of the potential threat of increases in wildfire frequency or severity to amphibians in the region. Incrementos Rápidos y Declinaciones Desfasadas en la Ocupación de Anfibios Después de un Incendio     

Trends and causes of severity, size, and number of fires in northwestern California, USA

Research in the last several years has indicated that fire size and frequency are on the rise in western U.S. forests. Although fire size and frequency are important, they do not necessarily scale with ecosystem effects of fire, as different ecosystems have different ecological and evolutionary relationships with fire. Our study assessed trends and patterns in fire size and frequency from 1910 to 2008 (all fires > 40 ha), and the percentage of high-severity in fires from 1987 to 2008 (all fires > 400 ha) on the four national forests of northwestern California. During 1910-2008, mean and maximum fire size and total annual area burned increased, but we found no temporal trend in the percentage of high-severity fire during 1987-2008. The time series of severity data was strongly influenced by four years with region-wide lightning events that burned huge areas at primarily low-moderate severity. Regional fire rotation reached a high of 974 years in 1984 and fell to 95 years by 2008. The percentage of high-severity fire in conifer-dominated forests was generally higher in areas dominated by smaller-diameter trees than in areas with larger-diameter trees. For Douglas-fir forests, the percentage of high-severity fire did not differ significantly between areas that re-burned and areas that only burned once (10% vs. 9%) when re-burned within 30 years. Percentage of high-severity fire decreased to 5% when intervals between first and second fires were > 30 years. In contrast, in both mixed-conifer and fir/high-elevation conifer forests, the percentage of high-severity fire was less when re-burned within 30 years compared to first-time burned (12% vs. 16% for mixed conifer; 11% vs. 19% for fir/high-elevation conifer). Additionally, the percentage of high-severity fire did not differ whether the re-burn interval was less than or greater than 30 years. Years with larger fires and greatest area burned were produced by region-wide lightning events, and characterized by less winter and spring precipitation than years dominated by smaller human-ignited fires. Overall percentage of high-severity fire was generally less in years characterized by these region-wide lightning events. Our results suggest that, under certain conditions, wildfires could be more extensively used to achieve ecological and management objectives in northwestern California.     

Effects of Postfire Salvage Logging on Deadwood‐Associated Beetles

Abstract: In Canada and the United States pressure to recoup financial costs of wildfire by harvesting burned timber is increasing, despite insufficient understanding of the ecological consequences of postfire salvage logging. We compared the species richness and composition of deadwood‐associated beetle assemblages among undisturbed, recently burned, logged, and salvage‐logged, boreal, mixed‐wood stands. Species richness was lowest in salvage‐logged stands, largely due to a negative effect of harvesting on the occurrence of wood‐ and bark‐boring species. In comparison with undisturbed stands, the combination of wildfire and logging in salvage‐logged stands had a greater effect on species composition than either disturbance alone. Strong differences in species composition among stand treatments were linked to differences in quantity and quality (e.g., decay stage) of coarse woody debris. We found that the effects of wildfire and logging on deadwood‐associated beetles were synergistic, such that the effects of postfire salvage logging could not be predicted reliably on the basis of data on either disturbance alone. Thus, increases in salvage logging of burned forests may have serious negative consequences for deadwood‐associated beetles and their ecological functions in early postfire successional forests.     

Postfire Logging in Riparian Areas

Abstract:  We reviewed the behavior of wildfire in riparian zones, primarily in the western United States, and the potential ecological consequences of postfire logging. Fire behavior in riparian zones is complex, but many aquatic and riparian organisms exhibit a suite of adaptations that allow relatively rapid recovery after fire. Unless constrained by other factors, fish tend to rebound relatively quickly, usually within a decade after a wildfire. Additionally, fire and subsequent erosion events contribute wood and coarse sediment that can create and maintain productive aquatic habitats over time. The potential effects of postfire logging in riparian areas depend on the landscape context and disturbance history of a site; however, available evidence suggests two key management implications: (1) fire in riparian areas creates conditions that may not require intervention to sustain the long-term productivity of the aquatic network and (2) protection of burned riparian areas gives priority to what is left rather than what is removed. Research is needed to determine how postfire logging in riparian areas has affected the spread of invasive species and the vulnerability of upland forests to insect and disease outbreaks and how postfire logging will affect the frequency and behavior of future fires. The effectiveness of using postfire logging to restore desired riparian structure and function is therefore unproven, but such projects are gaining interest with the departure of forest conditions from those that existed prior to timber harvest, fire suppression, and climate change. In the absence of reliable information about the potential consequence of postfire timber harvest, we conclude that providing postfire riparian zones with the same environmental protections they received before they burned is justified ecologically. Without a commitment to monitor management experiments, the effects of postfire riparian logging will remain unknown and highly contentious.     

Reconciling Salvage Logging of Boreal Forests with a Natural-Disturbance Management Model

Abstract:  In North American boreal forests, wildfire is the dominant agent of natural disturbance. A natural-disturbance model has therefore been promoted as an ecologically based approach to forest harvesting in these systems. Given accelerating resource demands, fire competes with harvest for timber, and there is increasing pressure to salvage naturally burned areas. This creates a management paradox: simultaneous promotion of natural disturbance as a guide to sustainability while salvaging forests that have been naturally disturbed. The major drivers of postfire salvage in Canadian boreal forests are societal perceptions, overallocation of forest resources, and economic and policy incentives, and postfire salvage compromises forest sustainability by diminishing the role of fire as a critical, natural process. These factors might be reconciled through consideration of fire in resource allocations and application of active adaptive management. We provide novel treatment of the role of burn severity in mediating biotic response by examining its influence on the amount, type, and distribution of live, postfire residual material, and we highlight the role of fire in shaping spatial and temporal patterns in forest biodiversity. Maintenance of natural postfire forests is a critical component of an ecosystem-based approach to forest management in boreal systems. Nevertheless, present practices focus heavily on expediting removal of timber from burned forests, despite increasing evidence that postfire communities differ markedly from postharvest systems, and there is a mismatch between emerging management models and past management practices. Policies that recognize the critical role of fire in these systems and facilitate enhanced understanding of natural system dynamics in support of development of sustainable management practices are urgently needed.     

Comparing modern and presettlement forest dynamics of a subboreal wilderness: does spruce budworm enhance fire risk?

Insect disturbance is often thought to increase fire risk through enhanced fuel loadings, particularly in coniferous forest ecosystems. Yet insect disturbances also affect successional pathways and landscape structure that interact with fire disturbances (and vice-versa) over longer time scales. We applied a landscape succession and disturbance model (LANDIS-II) to evaluate the relative strength of interactions between spruce budworm (Choristoneura fumiferana) outbreaks and fire disturbances in the Boundary Waters Canoe Area (BWCA) in northern Minnesota (USA). Disturbance interactions were evaluated for two different scenarios: presettlement forests and fire regimes vs. contemporary forests and fire regimes. Forest composition under the contemporary scenario trended toward mixtures of deciduous species (primarily Betula papyrifera and Populus spp.) and shade-tolerant conifers (Picea mariana, Abies balsamea, Thuja occidentalis), with disturbances dominated by a combination of budworm defoliation and high-severity fires. The presettlement scenario retained comparatively more "big pines" (i.e., Pinus strobus, P. resinosa) and tamarack (L. laricina), and experienced less budworm disturbance and a comparatively less-severe fire regime. Spruce budworm disturbance decreased area burned and fire severity under both scenarios when averaged across the entire 300-year simulations. Contrary to past research, area burned and fire severity during outbreak decades were each similar to that observed in non-outbreak decades. Our analyses suggest budworm disturbances within forests of the BWCA have a comparatively weak effect on long-term forest composition due to a combination of characteristics. These include strict host specificity, fine-scaled patchiness created by defoliation damage, and advance regeneration of its primary host, balsam fir (A. balsamea) that allows its host to persist despite repeated disturbances. Understanding the nature of the three-way interaction between budworm, fire, and composition has important ramifications for both fire mitigation strategies and ecosystem restoration initiatives. We conclude that budworm disturbance can partially mitigate long-term future fire risk by periodically reducing live ladder fuel within the mixed forest types of the BWCA but will do little to reverse the compositional trends caused in part by reduced fire rotations.     

Wildfire, Fuel Reduction, and Herpetofaunas across Diverse Landscape Mosaics in Northwestern Forests

Abstract:  The herpetofauna (amphibians and reptiles) of northwestern forests (U.S.A.) is diverse, and many species are locally abundant. Most forest amphibians west of the Cascade Mountain crest are associated with cool, cascading streams or coarse woody material on the forest floor, which are characteristics of mature forests. Extensive loss and fragmentation of habitat resulted from logging across approximately 50% of old-growth forests in northern California and approximately 80% of stands in Oregon and Washington. There is a complex landscape mosaic and overlap of northern and southern biotic elements in the Klamath-Siskiyou Region along the Oregon and California border, creating a biodiversity hotspot. The region experiences many low-severity fires annually, punctuated by periodic major fires, including the Biscuit fire, the largest in North America in 2002. In the fire's northern portion, severe fire occurred on >50% of stands of young, managed trees but on only about 25–33% of old-growth stands. This suggests that the legacy of timber harvest may produce fire-prone stands. Calls for prescribed fire and thinning to reduce fuel loads will remove large amounts of coarse woody material from forests, which reduces cover for amphibians and alters nutrient inputs to streams. Our preliminary evidence suggests no negative effects of wildfire on terrestrial amphibians, but stream amphibians decrease following wildfire. Most reptiles are adapted to open terrain, so fire usually improves their habitat. Today, the challenge is to maintain biodiversity in western forests in the face of intense political pressures designed to "prevent" catastrophic fires. We need a dedicated research effort to understanding how fire affects biota and to proactively investigate outcomes of fuel-reduction management on wildlife in western forests.     

Fire history and tree recruitment in the Colorado Front Range upper montane zone: implications for forest restoration

Forests experiencing moderate- or mixed-severity fire regimes are presumed to be widespread across the western United States, but few studies have characterized these complex disturbance regimes and their effects on contemporary forest structure. Restoration of pre-fire-suppression open-forest structure to reduce the risk of uncharacteristic stand-replacing fires is a guiding principle in forest management policy, but identifying which forests are clear candidates for restoration remains a challenge. We conducted dendroecological reconstructions of fire history and stand structure at 40 sites in the upper montane zone of the Colorado Front Range (2400-2800 m), sampled in proportion to the distribution of forest types in that zone (50% dominated by ponderosa pine, 28% by lodgepole pine, 12% by aspen, 10% by Douglas-fir). We characterized past fire severity based on remnant criteria at each site in order to assess the effect of fire history on tree establishment patterns, and we also evaluated the influence of fire suppression and climate. We found that 62% of the sites experienced predominantly moderate-severity fire, 38% burned at high severity, and no sites burned exclusively at low severity. The proportion of total tree and sapling establishment was significantly different among equal time periods based on a chi-square test, with highest tree and sapling establishment during the pre-fire-suppression period (1835-1919). Superposed epoch analysis revealed that fires burned during years of extreme drought (95% CI). The major pulse of tree establishment in the upper montane zone occurred during a multidecadal period of extreme drought conditions in the Colorado Front Range (1850-1889), during which 53% of the fires from the 1750-1989 period burned. In the upper montane zone of the Colorado Front Range, historical evidence suggests that these forests are resilient to prolonged periods of severe drought and associated severe fires.     

Forest productivity decline caused by successional paludification of boreal soils.

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

Temporal variability of forest fires in eastern Amazonia

Widespread occurrence of fires in Amazonian forests is known to be associated with extreme droughts, but historical data on the location and extent of forest fires are fundamental to determining the degree to which climate conditions and droughts have affected fire occurrence in the region. We used remote sensing to derive a 23-year time series of annual landscape-level burn scars in a fragmented forest of the eastern Amazon. Our burn scar data set is based on a new routine developed for the Carnegie Landsat Analysis System (CLAS), called CLAS-BURN, to calculate a physically based burn scar index (BSI) with an overall accuracy of 93% (Kappa coefficient 0.84). This index uses sub-pixel cover fractions of photosynthetic vegetation, non-photosynthetic vegetation, and shade/burn scar spectral end members. From 23 consecutive Landsat images processed with the CLAS-BURN algorithm, we quantified fire frequencies, the variation in fire return intervals, and rates of conversion of burned forest to other land uses in a 32 400 km2 area. From 1983 to 2007, 15% of the forest burned; 38% of these burned forests were subsequently deforested, representing 19% of the area cleared during the period of observation. While 72% of the fire-affected forest burned only once during the 23-year study period, 20% burned twice, 6% burned three times, and 2% burned four or more times, with the maximum of seven times. These frequencies suggest that the current fire return interval is 5-11 times more frequent than the estimated natural fire regime. Our results also quantify the substantial influence of climate and extreme droughts caused by a strong El Ni?o Southern Oscillation (ENSO) on the extent and likelihood of returning forest fires mainly in fragmented landscapes. These results are an important indication of the role of future warmer climate and deforestation in enhancing emissions from more frequently burned forests in the Amazon.     

Postfire response of flood-regenerating riparian vegetation in a semi-arid landscape

Piles of large wood (LW) deposited by major floods in river corridors can interact with naturally occurring wildfires from uplands to impact the regeneration of riparian vegetation. This study examines the spatial and short-term temporal response of riparian vegetation and soil nutrients to fire along the Sabie River, South Africa, with special emphasis on the effects of burned LW piles. At the study site there were 112 species of plants recorded with 28% of species restricted to the burned plots. As expected, vegetation cover was significantly lower in burned plots as compared with the unburned plots 12 months postfire. There was a significant influence of LW on species richness with fewer species recorded in the LW plots. For both fire and LW treatments, plant cover showed a significant change over three years. After an initial increase from 12 to 24 months (postfire) there was a decline in plant cover after 36 months. Species community composition was distinctly different between burned and unburned plots 12 months postfire, and the presence of LW affected species composition for burned plots but not for unburned ones. Time series ordination of LW plots highlighted the changes in species composition over the three years of sampling. Of trees with accumulations of LW within 5 m of their base, 48% had been killed by fire as compared to only 4% with no LW accumulations in close proximity. Soil-available P was significantly higher in the burned plots and even higher with burned LW while there were no effects on soil total N. There was also a significant positive trend between available P in soils and plant vegetation cover. Soil-exchangeable K was also significantly higher and total C significantly lower in the burned and LW plots. Burned plots also had significantly higher soil electrical conductivity (EC) and soil pH. The patchy nature of the studied fire, whose complexity is exacerbated by the distribution of flood deposited LW, acted to create a mosaic of alternate successional states as the riparian community recovers from flooding and the subsequent fire. We suspect that the resultant heterogeneity will increase ecosystem resilience by providing flexibility in the form of more options for a system response to subsequent disturbances.     

Climate effects on fire regimes and tree recruitment in Black Hills ponderosa pine forests

Climate influences forest structure through effects on both species demography (recruitment and mortality) and disturbance regimes. Here, I compare multi-century chronologies of regional fire years and tree recruitment from ponderosa pine forests in the Black Hills of southwestern South Dakota and northeastern Wyoming to reconstructions of precipitation and global circulation indices. Regional fire years were affected by droughts and variations in both Pacific and Atlantic sea surface temperatures. Fires were synchronous with La Ni?as, cool phases of the Pacific Decadal Oscillation (PDO), and warm phases of the Atlantic Multidecadal Oscillation (AMO). These quasi-periodic circulation features are associated with drought conditions over much of the western United States. The opposite pattern (El Ni?o, warm PDO, cool AMO) was associated with fewer fires than expected. Regional tree recruitment largely occurred during wet periods in precipitation reconstructions, with the most abundant recruitment coeval with an extended pluvial from the late 1700s to early 1800s. Widespread even-aged cohorts likely were not the result of large crown fires causing overstory mortality, but rather were caused by optimal climate conditions that contributed to synchronous regional recruitment and longer intervals between surface fires. Synchronous recruitment driven by climate is an example of the Moran effect. The presence of abundant fire-scarred trees in multi-aged stands supports a prevailing historical model for ponderosa pine forests in which recurrent surface fires affected heterogenous forest structure, although the Black Hills apparently had a greater range of fire behavior and resulting forest structure over multi-decadal time scales than ponderosa pine forests of the Southwest that burned more often.     

Small-scale fuel variation alters fire intensity and shrub abundance in a pine savanna

Small-scale variation in fire intensity and effects may be an important source of environmental heterogeneity in frequently burned plant communities. We hypothesized that variation in fire intensity resulting from local differences in fuel loads produces heterogeneity in pine savanna ground cover by altering shrub abundance. To test this hypothesis, we experimentally manipulated prefire fuel loads to mimic naturally occurring fuel-load heterogeneity associated with branch falls, needle fall near large pines, and animal disturbances in a frequently burned longleaf pine (Pinus palustris) savanna in Louisiana, USA. We applied one of four fuel treatments (unaltered control, fine-fuel removal, fine-fuel addition, wood addition) to each of 540 (1-m2) quadrats prior to growing-season prescribed fires in each of two years (1999 and 2001). In both years fuel addition increased (and fuel removal decreased) fuel consumption and maximum fire temperatures relative to unaltered controls. Fuel addition, particularly wood, increased damage to shrubs, increased shrub mortality, and decreased resprout density relative to controls. We propose that local variation in fire intensity may contribute to maintenance of high species diversity in pine savannas by reducing shrub abundance and creating openings in an otherwise continuous ground cover.     

Avifaunal responses to fire in southwestern montane forests along a burn severity gradient.

The effects of burn severity on avian communities are poorly understood, yet this information is crucial to fire management programs. To quantify avian response patterns along a burn severity gradient, we sampled 49 random plots (2001-2002) at the 17351-ha Cerro Grande Fire (2000) in New Mexico, USA. Additionally, pre-fire avian surveys (1986-1988, 1990) created a unique opportunity to quantify avifaunal changes in 13 pre-fire transects (resampled in 2002) and to compare two designs for analyzing the effects of unplanned disturbances: after-only analysis and before-after comparisons. Distance analysis was used to calculate densities. We analyzed after-only densities for 21 species using gradient analysis, which detected a broad range of responses to increasing burn severity: (I) large significant declines, (II) weak, but significant declines, (III) no significant density changes, (IV) peak densities in low- or moderate-severity patches, (V) weak, but significant increases, and (VI) large significant increases. Overall, 71% of the species included in the after-only gradient analysis exhibited either positive or neutral density responses to fire effects across all or portions of the severity gradient (responses III-VI). We used pre/post pairs analysis to quantify density changes for 15 species using before-after comparisons; spatiotemporal variation in densities was large and confounded fire effects for most species. Only four species demonstrated significant effects of burn severity, and their densities were all higher in burned compared to unburned forests. Pre- and post-fire community similarity was high except in high-severity areas. Species richness was similar pre- and post-fire across all burn severities. Thus, ecosystem restoration programs based on the assumption that recent severe fires in Southwestern ponderosa pine forests have overriding negative ecological effects are not supported by our study of post-fire avian communities. This study illustrates the importance of quantifying burn severity and controlling confounding sources of spatiotemporal variation in studies of fire effects. After-only gradient analysis can be an efficient tool for quantifying fire effects. This analysis can also augment historical data sets that have small samples sizes coupled with high non-process variation, which limits the power of before-after comparisons.     

Frequent fire alters nitrogen transformations in ponderosa pine stands of the inland northwest

Recurrent, low-severity fire in ponderosa pine (Pinus ponderosa)/interior Douglas-fir (Pseudotsuga menziesii var. glauca) forests is thought to have directly influenced nitrogen (N) cycling and availability. However, no studies to date have investigated the influence of natural fire intervals on soil processes in undisturbed forests, thereby limiting our ability to understand ecological processes and successional dynamics in this important ecosystem of the Rocky Mountain West. Here, we tested the standing hypothesis that recurrent fire in ponderosa pine/Douglas-fir forests of the Inland Northwest decreases total soil N, but increases N turnover and nutrient availability. We compared soils in stands unburned over the past 69-130 years vs. stands exposed to two or more fires over the last 130 years at seven distinct locations in two wilderness areas. Mineral soil samples were collected from each of the seven sites in June and July of 2003 and analyzed for pH, total C and N, potentially mineralizable N (PMN), and extractable NH4+, NO3-, PO4(-3), Ca+2, Mg+2, and K+. Nitrogen transformations were assessed at five sites by installing ionic resin capsules in the mineral soil in August of 2003 and by conducting laboratory assays of nitrification potential and net nitrification in aerobic incubations. Total N and PMN decreased in stands subjected to multiple fires. This loss of total N and labile N was not reflected in concentrations of extractable NH4+ and NO3-. Rather, multiple fires caused an increase in NO3 sorbed on ionic resins, nitrification potential, and net nitrification in spite of the burned stands not having been exposed to fire for at least 12-17 years. Charcoal collected from a recent fire site and added to unburned soils increased nitrification potential, suggesting that the decrease of charcoal in the absence of fire may play an important role in N transformations in fire-dependent ecosystems in the long term. Interestingly, we found no consistent effect of fire frequency on extractable P or alkaline metal concentrations. Our results corroborate the largely untested hypothesis that frequent fire in ponderosa pine forests increases inorganic N availability in the long term and emphasize the need to study natural, unmanaged sites in far greater detail.     

青藏高原东部亚高山森林草甸植被地理格局的成因探讨

森林和草地在亚高山带的阴阳坡上形成的交错分布现象,呈一弧型分布在青藏高原东部本文在综合分析了造成这种现象的生理生态因素之后,重点从火的作用、放牧作用和历史时期人类活动的影响等方面,讨论了青藏高原东部亚高山带植被地理格局的形成原因结果表明,自然生态因素和人类活动的影响在植被地理格局的形成过程中起着同等重要的作用,其中火灾的作用以及气候变化造成的人口迁移压力作用十分明显．在这一弧形区域内,林线以下的亚高山灌丛或草甸多属次生类型,人类有目的的烧荒和长期放牧阻滞了植被的顺向演替,维持了亚高山（灌丛）草甸的偏途顶级状态．