Effect of prior disturbances on the extent and severity of wildfire in Colorado subalpine forests

Disturbances are important in creating spatial heterogeneity of vegetation patterns that in turn may affect the spread and severity of subsequent disturbances. Between 1997 and 2002 extensive areas of subalpine forests in northwestern Colorado were affected by a blowdown of trees, bark beetle outbreaks, and salvage logging. Some of these stands were also affected by severe fires in the late 19th century. During a severe drought in 2002, fires affected extensive areas of these subalpine forests. We evaluated and modeled the extent and severity of the 2002 fires in relation to these disturbances that occurred over the five years prior to the fires and in relation to late 19th century stand-replacing fires. Occurrence of disturbances prior to 2002 was reconstructed using a combination of tree-ring methods, aerial photograph interpretation, field surveys, and geographic information systems (GIS). The extent and severity of the 2002 fires were based on the normalized difference burn ratio (NDBR) derived from satellite imagery. GIS and classification trees were used to analyze the effects of prefire conditions on the 2002 fires. Previous disturbance history had a significant influence on the severity of the 2002 fires. Stands that were severely blown down (> 66% trees down) in 1997 burned more severely than other stands, and young (approximately 120 year old) postfire stands burned less severely than older stands. In contrast, prefire disturbances were poor predictors of fire extent, except that young (approximately 120 years old) postfire stands were less extensively burned than older stands. Salvage logging and bark beetle outbreaks that followed the 1997 blowdown (within the blowdown as well as in adjacent forest that was not blown down) did not appear to affect fire extent or severity. Conclusions regarding the influence of the beetle outbreaks on fire extent and severity are limited, however, by spatial and temporal limitations associated with aerial detection surveys of beetle activity. Thus, fire extent in these forests is largely independent of prefire disturbance history and vegetation conditions. In contrast, fire severity, even during extreme fire weather and in conjunction with a multiyear drought, is influenced by prefire stand conditions, including the history of previous disturbances.     

Demography of northern flying squirrels informs ecosystem management of western interior forests.

We studied northern flying squirrel (Glaucomys sabrinus) demography in the eastern Washington Cascade Range to test hypotheses about regional and local abundance patterns and to inform managers of the possible effects of fire and fuels management on flying squirrels. We quantified habitat characteristics and squirrel density, population trends, and demography in three typical forest cover types over a four-year period. We had 2034 captures of flying squirrels over 41 000 trap nights from 1997 through 2000 and marked 879 squirrels for mark-recapture population analysis. Ponderosa pine (Pinus ponderosa) forest appeared to be poorer habitat for flying squirrels than young or mature mixed-conifer forest. About 35% fewer individuals were captured in open pine forest than in dry mixed-conifer Douglas-fir (Pseudotsuga menziesii) and grand fir (Abies grandis) forests. Home ranges were 85% larger in pine forest (4.6 ha) than in mixed-conifer forests (2.5 ha). Similarly, population density (Huggins estimator) in ponderosa pine forest was half (1.1 squirrels/ha) that of mixed-conifer forest (2.2 squirrels/ha). Tree canopy cover was the single best correlate of squirrel density (r = 0.77), with an apparent threshold of 55% canopy cover separating stands with low- from high-density populations. Pradel estimates of annual recruitment were lower in open pine (0.28) than in young (0.35) and mature (0.37) forest. High recruitment was most strongly associated with high understory plant species richness and truffle biomass. Annual survival rates ranged from 45% to 59% and did not vary among cover types. Survival was most strongly associated with understory species richness and forage lichen biomass. Maximum snow depth had a strong negative effect on survival. Rate of per capita increase showed a density-dependent response. Thinning and prescribed burning in ponderosa pine and dry mixed conifer forests to restore stable fire regimes and forest structure might reduce flying squirrel densities at stand levels by reducing forest canopy, woody debris, and the diversity or biomass of understory plants, truffles, and lichens. Those impacts might be ameliorated by patchy harvesting and the retention of large trees, woody debris, and mistletoe brooms. Negative stand-level impacts would be traded for increased resistance and resilience of dry-forest landscapes to now-common, large-scale stand replacement fires.     

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.     

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.     

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.     

The Effects of Blister Rust on Post-Fire Regeneration of Whitebark Pine: The Sundance Burn of Northern Idaho (U.S.A.)

In the northern Rocky Mountains, whitebark pine ( Pinus albicaulis ) is rapidly declining as a result of previous fire exclusion policies, mountain pine beetle ( Dendroctonus ponderosae ) outbreaks, and white pine blister rust ( Cronartium ribicola ). Blister rust is potentially the most destructive agent, killing seedlings, cone-bearing branches, and, eventually, mature trees. We examined densities of whitebark pine regeneration and the incidence and severity of blister-rust infection of seedlings and saplings in the 25-year-old Sundance Burn in the Selkirk Range of northern Idaho, an area heavily infected by blister rust. We found that the mean regeneration density of whitebark pine was significantly lower than that of two other comparably aged burns in western Montana. The low density was attributed to the severe damage to the seed source on the burn perimeter, resulting from previous infestation of mountain pine beetle and blister rust. Overall, 29% of the whitebark pine regeneration in the Sundance Burn was infected by blister rust. Age and height of seedlings were important predictors of incidence of infection, and height was the most important predictor of severity of infection. Thus, as seedlings grow larger, they present a bigger target to airborne blister-rust spores. Because of the lack of seed production in the adjacent forest and expected mortality, regeneration of whitebark pine in the Sundance Burn will be slow. In areas of northern Idaho and northwestern Montana affected by blister rust and pine beetle, prescribed fires for managing whitebark pine ecosystems should be restricted to small areas or should require plantings of rust-resistant seedlings.     

Meta-analysis of avian and small-mammal response to fire severity and fire surrogate treatments in U.S. fire-prone forests

Management in fire-prone ecosystems relies widely upon application of prescribed fire and/or fire surrogate (e.g., forest thinning) treatments to maintain biodiversity and ecosystem function. Recently, published literature examining wildlife response to fire and fire management has increased rapidly. However, none of this literature has been synthesized quantitatively, precluding assessment of consistent patterns of wildlife response among treatment types. Using meta-analysis, we examined the scientific literature on vertebrate demographic responses to burn severity (low/moderate, high), fire surrogates (forest thinning), and fire and fire surrogate combined treatments in the most extensively studied fire-prone, forested biome (forests of the United States). Effect sizes (magnitude of response) and their 95% confidence limits (response consistency) were estimated for each species-by-treatment combination with two or more observations. We found 41 studies of 119 bird and 17 small-mammal species that examined short-term responses (< or =4 years) to thinning, low/moderate- and high-severity fire, and thinning plus prescribed fire; data on other taxa and at longer time scales were too sparse to permit quantitative assessment. At the stand scale (<50 ha), thinning and low/moderate-severity fire demonstrated similar response patterns in these forests. Combined thinning plus prescribed fire produced a higher percentage of positive responses. High-severity fire provoked stronger responses, with a majority of species possessing higher or lower effect sizes relative to fires of lower severity. In the short term and at fine spatial scales, fire surrogate forest-thinning treatments appear to effectively mimic low/moderate-severity fire, whereas low/moderate-severity fire is not a substitute for high-severity fire. The varied response of taxa to each of the four conditions considered makes it clear that the full range of fire-based disturbances (or their surrogates) is necessary to maintain a full complement of vertebrate species, including fire-sensitive taxa. This is especially true for high-severity fire, where positive responses from many avian taxa suggest that this disturbance (either as wildfire or prescribed fire) should be included in management plans where it is consistent with historic fire regimes and where maintenance of regional vertebrate biodiversity is a goal.     

A diameter increment model for Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) stands in Isparta region, Turkey

The purpose of this research was to test the precision of a diameter increment model for the estimation of future periodic diameter increment. Individual trees of Crimean pine (Pinus nigra Arnold) and Calabrian pine (Pinus brutia Ten.) located in both natural and plantation stands were selected. For that reason, normal closed canopy, pure, even-aged and undisturbed stands were examined. In 2002, plots were sampled in three natural and three plantation stands in Isparta region of Turkey. The number of sampling points in sample plots ranged from 19 to 55. In each sampling point, a subject tree and six competitors were selected. In each sampling point, subject tree and competitor trees were stem mapped (x and y coordinate system), and diameter (dbh), total height, age, and 10-yrs radial increment recorded. The predictors of a distance dependent diameter increment model were chosen that included tree level (diameter (d), competition index (CI), and age (t)) and stand level (basal area (G), and site index (SI)) characteristics as well as their transformations. The best fit index of the regression model was pursued in trials with variable combinations. The models explained 65%, 60%, 68% and 50% of the variation in individual tree diameter increment of Crimean pine and Calabrian pine for both natural and plantations stands, respectively. These models can be estimated diameter increment of individual trees at highly significant level (p<0.001).     

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.     

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.     

Multiple disturbances accelerate clonal growth in a potentially monodominant bamboo

Organisms capable of rapid clonal growth sometimes monopolize newly freed space and resources. We hypothesize that sequential disturbances might change short-term clonal demography of these organisms in ways that promote formation of monotypic stands. We examined this hypothesis by studying the clonal response of Arundinaria gigantea (giant cane, a bamboo) to windstorm and fire. We studied giant cane growing in both a large tornado-blowdown gap and under forest canopy, in burned and unburned plots, using a split-block design. We measured density of giant cane ramets (culms) and calculated finite rates of increase (lamda) for populations of ramets over three years. Ramet density nearly doubled in stands subjected to both windstorm and fire; the high ramet densities that resulted could inhibit growth in other plants. In comparison, ramet density increased more slowly after windstorm alone, decreased after fire alone, and remained in stasis in controls. We predict that small, sparse stands of giant cane could spread and amalgamate to form dense, monotypic stands (called "canebrakes") that might influence fire return intervals and act as an alternative state to bottomland forest. Other clonal species may similarly form monotypic stands following successive disturbances via rapid clonal growth.     

Forest structure and light regimes following moderate wind storms: implications for multi-cohort management.

Moderate-severity disturbances appear to be common throughout much of North America, but they have received relatively little detailed study compared to catastrophic disturbances and small gap dynamics. In this study, we examined the immediate impact of moderate-intensity wind storms on stand structure, opening sizes, and light regimes in three hemlock-hardwood forests of northeastern Wisconsin. These were compared to three stands managed by single-tree and group selection, the predominant forest management system for northern hardwoods in the region. Wind storms removed an average of 41% of the stand basal area, compared to 27% removed by uneven-aged harvests, but both disturbances removed trees from a wide range of size classes. The removal of nearly half of the large trees by wind in two old-growth stands caused partial retrogression to mature forest structure, which has been hypothesized to be a major disturbance pathway in the region. Wind storms resulted in residual stand conditions that were much more heterogeneous than in managed stands. Gap sizes ranged from less than 10 m2 up to 5000 m2 in wind-disturbed stands, whereas the largest opening observed in managed stands was only 200 m2. Wind-disturbed stands had, on average, double the available solar radiation at the forest floor compared to managed stands. Solar radiation levels were also more heterogeneous in wind-disturbed stands, with six times more variability at small scales (0.1225 ha) and 15 times more variability at the whole-stand level. Modification of uneven-aged management regimes to include occasional harvests of variable intensity and spatial pattern may help avoid the decline in species diversity that tends to occur after many decades of conventional uneven-aged management. At the same time, a multi-cohort system with these properties would retain a high degree of average crown cover, promote structural heterogeneity typical of old-growth forests, and maintain dominance by late-successional species.     

Seed predation and selection exerted by a seed predator influence subalpine tree densities

Strongly interacting species often have pronounced direct and indirect effects on other species. Here we focus of the effects of pine squirrels (Tamiasciurus spp.), which are a dominant pre-dispersal seed predator of many conifers including limber pines (Pinus flexilis) and whitebark pines (P. albicaulis). Pine squirrels depress seed abundance by harvesting most limber and whitebark pine cones on their territories. Pine squirrels further reduce seed availability for Clark's Nutcrackers (Nucifraga columbiana), the primary seed disperser of these pines, because selection exerted by pine squirrels has reduced the number of seeds per cone and causes seeds to be less accessible. We predicted that, if fewer seeds were available for dispersal by nutcrackers, pine recruitment should be suppressed in areas with pine squirrels. In support of this prediction, stand densities were about two times greater in areas where pine squirrels are absent than in areas where they are present. Alternative explanations that we considered do not account for these differences; however, precipitation may limit stand densities in the absence of seed limitation by pine squirrels. In sum, pine squirrels apparently depress limber and whitebark pine stand densities, with the potential for ecosystem impacts because these pines are foundation species within Western subalpine ecosystems.     

Corridors promote fire via connectivity and edge effects

Landscape corridors, strips of habitat that connect otherwise isolated habitat patches, are commonly employed during management of fragmented landscapes. To date, most reported effects of corridors have been positive; however, there are long-standing concerns that corridors may have unintended consequences. Here, we address concerns over whether corridors promote propagation of disturbances such as fire. We collected data during prescribed fires in the world's largest and best replicated corridor experiment (Savannah River Site, South Carolina, USA), six -50-ha landscapes of open (shrubby/herbaceous) habitat within a pine plantation matrix, to test several mechanisms for how corridors might influence fire. Corridors altered patterns of fire temperature through a direct connectivity effect and an indirect edge effect. The connectivity effect was independent of fuel levels and was consistent with a hypothesized wind-driven "bellows effect." Edges, a consequence of corridor implementation, elevated leaf litter (fuel) input from matrix pine trees, which in turn increased fire temperatures. We found no evidence for corridors or edges impacting patterns of fire spread: plots across all landscape positions burned with similar probability. Impacts of edges and connectivity on fire temperature led to changes in vegetation: hotter-burning plots supported higher bunch grass cover during the field season after burning, suggesting implications for woody/herbaceous species coexistence. To our knowledge, this represents the first experimental evidence that corridors can modify landscape-scale patterns of fire intensity. Corridor impacts on fire should be carefully considered during landscape management, both in the context of how corridors connect or break distributions of fuels and the desired role of fire as a disturbance, which may range from a management tool to an agent to be suppressed. In our focal ecosystem, longleaf pine woodland, corridors might provide a previously unrecognized benefit during prescribed burning activities, by promoting fire intensity, which may assist in promoting plant biodiversity.     

Thinning and burning result in low-level invasion by nonnative plants but neutral effects on natives.

Many historically fire-adapted forests are now highly susceptible to damage from insects, pathogens, and stand-replacing fires. As a result, managers are employing treatments to reduce fuel loadings and to restore the structure, species, and processes that characterized these forests prior to widespread fire suppression, logging, and grazing. However, the consequences of these activities for understory plant communities are not well understood. We examined the effects of thinning and prescribed fire on plant composition and diversity in Pinus ponderosa forests of eastern Washington (USA). Data on abundance and richness of native and nonnative plants were collected in 70 stands in the Colville, Okanogan, and Wenatchee National Forests. Stands represented one of four treatments: thinning, burning, thinning followed by burning, or control; treatments had been conducted 3-19 years before sampling. Multi-response permutation procedures revealed no significant effect of thinning or burning on understory plant composition. Similarly, there were no significant differences among treatments in cover or richness of native plants. In contrast, nonnative plants showed small, but highly significant, increases in cover and richness in response to both thinning and burning. In the combined treatment, cover of nonnative plants averaged 2% (5% of total plant cover) but did not exceed 7% (16% of total cover) at any site. Cover and richness of nonnative herbs showed small increases with intensity of disturbance and time since treatment. Nonnative plants were significantly less abundant in treated stands than on adjacent roadsides or skid trails, and cover within these potential source areas explained little of the variation in abundance within treated stands. Although thinning and burning may promote invasion of nonnative plants in these forests, our data suggest that their abundance is limited and relatively stable on most sites.     

Pyric Herbivory: Rewilding Landscapes through the Recoupling of Fire and Grazing

Abstract:  Our understanding of fire and grazing is largely based on small-scale experimental studies in which treatments are uniformly applied to experimental units that are considered homogenous. Any discussion of an interaction between fire and grazing is usually based on a statistical approach that ignores the spatial and temporal interactions on complex landscapes. We propose a new focus on the ecological interaction of fire and grazing in which each disturbance is spatially and temporally dependent on the other and results in a landscape where disturbance is best described as a shifting mosaic (a landscape with patches that vary with time since disturbance) that is critical to ecological structure and function of many ecosystems. We call this spatiotemporal interaction pyric herbivory (literal interpretation means grazing driven by fire). Pyric herbivory is the spatial and temporal interaction of fire and grazing, where positive and negative feedbacks promote a shifting pattern of disturbance across the landscape. We present data we collected from the Tallgrass Prairie Preserve in the southern Great Plains of North America that demonstrates that the interaction between free-roaming bison ( Bison bison ) and random fires promotes heterogeneity and provides the foundation for biological diversity and ecosystem function of North American and African grasslands. This study is different from other studies of fire and grazing because the fires we examined were random and grazing animals were free to roam and select from burned and unburned patches. For ecosystems across the globe with a long history of fire and grazing, pyric herbivory with any grazing herbivore is likely more effective at restoring evolutionary disturbance patterns than a focus on restoring any large vertebrate while ignoring the interaction with fire and other disturbances .     

环境因子和火干扰因子对喀纳斯泰加林柳兰分布的影响

喀纳斯泰加林植被的生长和分布受火干扰和环境因子的共同作用,为了解喀纳斯泰加林林下柳兰分布与生态因子的关系,促进种群扩繁,共设置了35个样地进行调查和分析。建立柳兰重要值与生态因子矩阵,利用CCA分析法对不同火烧林分中柳兰的生长分布状况与林火烈度、火后时间、海拔、坡度、坡向、郁闭度、土壤pH值、速效钾、土壤有机质、灌木层盖度、草本层盖度和草本层物种多样性指数等20个生态因子的关系进行了分析。结果显示,第一轴主要反映林火烈度、海拔、坡向,灌木层盖度、pH值、有机质、全钾和速效钾的变化;第二轴主要反映林火烈度、火后时间、草本层盖度、坡向、坡位、土壤容重、电导率、有机质、有效磷和全氮的变化。对柳兰分布有显著影响的生态因子是林火烈度、有机质、海拔、坡向、灌木层盖度、土壤容重、全钾和速效钾,前两轴的累计贡献率为72.48%。柳兰适合在土壤呈弱酸性、海拔低、阳坡、土壤容重小、土壤有机质、全钾和速效钾含量高的环境中生长。柳兰是阳性植物,是火烧后的先锋植物。对影响柳兰分布的生态因子进行定量分离,结果发现环境因子和火干扰因子对柳兰样地物种分布的解释能力为88.2%,显示出较好的排序效果。喀纳斯泰加林受人为干扰较少,柳兰所在植物群落与环境保持了良好的对应关系,生态因子与物种分布呈显著相关(P=0.003),表明CCA排序结果可以解释生态因子对物种分布的影响程度。     

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.