Eight years of seasonal burning and herbicidal brush control influence sapling longleaf pine growth,understory vegetation,and the outcome of an ensuing wildfire

To study how fire or herbicide use influences longleaf pine (Pinus palustris Mill.) overstory and understory vegetation, five treatments were initiated in a 5–6-year-old longleaf pine stand: check, biennial arborescent plant control by directed herbicide application, and biennial burning in March, May, or July. The herbicide or prescribed fire treatments were applied in 1999, 2001, 2003, and 2005. All prescribed fires were intense and averaged 700 kJ/s/m of fire front across all 12 burns. Using pretreatment variables as covariates, longleaf pine survival and volume per hectare were significantly less on the three prescribed fire treatments than on checks. Least-square means in 2006 for survival were 70, 65, 64, 58, and 56% and volume per hectare was 129, 125, 65, 84, and 80 m³/ha on the check, herbicide, March-, May-, and July-burn treatments, respectively. A wildfire in March 2007 disproportionately killed pine trees on the study plots. In October 2007, pine volume per hectare was 85, 111, 68, 98, and 93 m³/ha and survival was 32, 41, 53, 57, and 55% on the check, herbicide, March-, May-, and July-burn treatments, respectively, after dropping trees that died through January 2009 from the database. Understory plant cover was also affected by treatment and the ensuing wildfire. In September 2006, herbaceous plant cover averaged 4% on the two unburned treatments and 42% on the three prescribed fire treatments. Seven months after the wildfire, herbaceous plant cover averaged 42% on the two previously unburned treatments and 50% on the three prescribed fire treatments. Before the wildfire, understory tree cover was significantly greater on checks (15%) than on the other four treatments (1.3%), but understory tree cover was similar across all five treatments 7 months after the wildfire averaging 1.1%. The greater apparent intensity of the wildfire on the previously unburned treatments most likely resulted from a greater accumulation of fuels on the check and herbicide plots that also collectively had a higher caloric content than fuels on the biennially prescribed burned plots. These results showed the destructive force of wildfire to overstory trees in unburned longleaf pine stands while also demonstrating the rejuvenating effects of wildfire within herbaceous plant communities. They caution for careful reintroduction of prescribed fire even if fire was excluded for less than a decade.     

The role of herbicide in savanna restoration: Effects of shrub reduction treatments on the understory and overstory of a longleaf pine flatwoods

Woody plant encroachment is a threat to savanna ecosystems worldwide. By exploiting differences in the physiology and seasonality of herbaceous species and encroaching hardwoods, herbicides can be used to control woody shrubs in savannas without causing lasting harm to desirable vegetation. We applied three herbicides and one tank mix to control shrubs following removal of the slash pine (Pinus elliottii Engelm.) canopy and replanting with container-grown longleaf pine (Pinus palustris Mill.) seedlings in a mesic-wet savanna in the southeastern USA. The herbicides tested were imazapyr, sulfometuron methyl, hexazinone, and a hexazinone + sulfometuron methyl tank mix. 4 years after application, no negative effects on understory species richness, diversity, evenness, or community composition were evident in any of the herbicide treatments. Oaks (Quercus spp.), one of the dominant shrub genera on the study site, were resistant to sulfometuron methyl, and this herbicide was therefore ineffective both as a pine release treatment and for enhancing herbaceous species cover. Imazapyr was the most effective treatment overall, leading to significant improvements in longleaf pine seedling growth and also enhancing herbaceous species cover. Both hexazinone and the hexazinone + sulfometuron methyl tank mix provided some seedling growth and understory enhancement as well. In particular, the tank mix significantly increased wiregrass cover relative to the control. Shrubs resprouted quickly following a dormant-season prescribed fire in the fifth year after treatment, indicating that herbicide-related increases in herbaceous cover may be lost if an aggressive prescribed fire program is not implemented.     

Survival and growth of upland oak and co-occurring competitor seedlings following single and repeated prescribed fires

Studies within and outside the U.S. indicate recurring oak (Quercus spp.) regeneration problems. In deciduous forests of the eastern U.S., a prevailing explanation for this trend is fire suppression leading to high competitor abundance and low understory light. In response, prescribed fire is increasingly used as a management tool to remedy these conditions and encourage future oak establishment and growth. Within eastern Kentucky, we implemented single and repeated (3×) prescribed fires over a 6-yr period (2002–2007). Pre- and post-burn, we quantified canopy cover and oak seedling survival and growth compared to other woody seedlings deemed potential competitors, primarily red maple (Acer rubrum L.) and sassafras (Sassafras albidum (Nutt.) Nees.). Burning temporarily decreased canopy cover 3–10%, but cover rebounded the subsequent growing season. Repeated burning ultimately produced canopy cover about 6% lower than sites unburned and burned once, suggesting a cumulative effect on understory light. Red maple exhibited low survival (∼40%) following single and repeated burns, but growth remained similar to unburned seedlings. Burning had little impact on sassafras survival and led to total height and basal diameters 2× greater than unburned seedlings. A single burn had no impact on red oak (Erythrobalanus spp.) survival and increased height and basal diameters 25–30%, but this positive growth response was driven by seedlings on several plots which experienced high burn temperatures and consequently high overstory mortality. White oaks (Leucobalanus spp.), however, exhibited twice as high mortality compared to those unburned, with no change in growth parameters. Repeated burning negatively impacted survival and growth of both oak groups compared to unburned seedlings. With both burn regimes, oaks with smaller pre-burn basal diameters exhibited the lowest post-burn survival. Thus, despite the ability of prescribed burns to temporarily increase understory light and reduce red maple survival, neither single or repeated burns placed oaks in an improved competitive position. These findings result from a combination of highly variable yet interdependent factors including the (1) life history traits of oaks compared to their co-occurring competitors, (2) pre-burn stature of pre-existing oak seedlings, and (3) variability in fire temperature and effects on understory light.     

Structure and composition changes following restoration treatments of longleaf pine forests on the Gulf Coastal Plain of Alabama

Longleaf pine (Pinus palustris Mill.) forests of the Gulf Coastal Plain historically burned every 2–4 years with low intensity fires, which maintained open stands with herbaceous dominated understories. During the early and mid 20th century however, reduced fire frequency allowed fuel to accumulate and hardwoods to increase in the midstory and overstory layers, while woody shrubs gained understory dominance. In 2001, a research study was installed in southern Alabama to develop management options that could be used to reduce fuel loads and restore the ecosystem. As part of a nationwide fire and fire surrogates study, treatments included a control (no fire or other disturbance), prescribed burning only, thinning of selected trees, thinning plus prescribed burning, and herbicide plus prescribed burning. After two cycles of prescribed burning, applied biennially during the growing season, there were positive changes in ecosystem composition. Although thinning treatments produced revenue, while reducing midstory hardwoods and encouraging growth of a grassy understory, burning was needed to discourage regrowth of the hardwood midstory and woody understory. Herbicide application followed by burning gave the quickest changes in understory composition, but repeated applications of fire eventually produced the same results at the end of this 8-year study. Burning was found to be a critical component of any restoration treatment for longleaf communities of this region with positive changes in overstory, midstory and understory layers after just three or four burns applied every 2 or 3 years.     

Understory vegetation response after 30 years of interval prescribed burning in two ponderosa pine sites in northern Arizona,USA

Southwestern USA ponderosa pine (Pinus ponderosa C. Lawson var. scopulorum Engelm.) forests evolved with frequent surface fires and have changed dramatically over the last century. Overstory tree density has sharply increased while abundance of understory vegetation has declined primarily due to the near cessation of fires. We examined effects of varying prescribed fire-return intervals (1, 2, 4, 6, 8, and 10 years, plus unburned) on the abundance and composition of understory vegetation in 2007 and 2008 after 30+ years of fall prescribed burning at two ponderosa pine sites. We found that after 30 years, overstory canopy cover remained high, while understory plant canopy cover was low, averaging <12% on all burn intervals. We attributed the weak understory response to a few factors – the most important of which was the high overstory cover at both sites. Graminoid cover and cover of the major grass species, Elymus elymoides (squirreltail), increased on shorter fire-return intervals compared to unburned plots, but only at one site. Community composition differed significantly between shorter fire-return intervals and unburned plots at one site, but not the other. For several response variables, precipitation levels appeared to have a stronger effect than treatments. Our findings suggest that low-severity burn treatments in southwestern ponderosa pine forests, especially those that do not decrease overstory cover, are minimally effective in increasing understory plant cover. Thinning of these dense forests along with prescribed burning is necessary to increase cover of understory vegetation.     

Thinning and chipping small-diameter ponderosa pine changes understory plant communities on the Colorado Front Range

Novel fire mitigation treatments that chip harvested biomass on site are increasingly prescribed to reduce the density of small-diameter trees, yet the ecological effects of these treatments are unknown. Our objective was to investigate the impacts of mechanical thinning and whole tree chipping on Pinus ponderosa (ponderosa pine) regeneration and understory plant communities to guide applications of these new fuel disposal methods. We sampled in three treatments: (1) unthinned forests (control), (2) thinned forests with harvested biomass removed (thin-only), and (3) thinned forests with harvested biomass chipped and broadcast on site (thin + chip). Plots were located in a ponderosa pine forest of Colorado and vegetation was sampled three to five growing seasons following treatment. Forest litter depth, augmented with chipped biomass, had a negative relationship with cover of understory plant species. In situ chipping often produces a mosaic of chipped patches tens of meters in size, creating a range of woodchip depths including areas lacking woodchip cover within thinned and chipped forest stands. Thin-only and thin + chip treatments had similar overall abundance and species richness of understory plants at the stand scale, but at smaller spatial scales, areas within thin + chip treatments that were free of woodchip cover had an increased abundance of understory vegetation compared to all other areas sampled. Relative cover of non-native plant species was significantly higher in the thin-only treatments compared to control and thin + chip areas. Thin + chip treated forests also had a significantly different understory plant community composition compared to control or thin-only treatments, including an increased richness of rhizomatous plant species. We suggest that thinning followed by either chipping or removing the harvested biomass could alter understory plant species composition in ponderosa pine forests of Colorado. When considering post-treatment responses, managers should be particularly aware of both the depth and the distribution of chipped biomass that is left in forested landscapes.     

A comparison of fire hazard mitigation alternatives in pinyon–juniper woodlands of Arizona

Concern over uncontrollable wildfire in pinyon–juniper woodlands has led public land managers in the southwestern United States to seek approaches for mitigating wildfire hazard, yet little information is available concerning effectiveness and ecological responses of alternative treatments. We established a randomized block experiment at a pinyon–juniper site in northern Arizona and tested effects of no treatment (Control), thinning only (Thin), prescribed fire only (Burn), and thinning followed by prescribed fire (Thin + Burn) on overstory structure, hazardous fuels reduction, and woody understory responses. One year after implementation, mean trees per hectare (TPH) of Utah juniper (Juniperus osteosperma) and pinyon pine (Pinus edulis), and basal area (BA) of pinyon, were significantly (P < 0.05) less in Thin and Thin + Burn treatments than Control. Additionally, pinyon TPH was less in Burn than Control. Quadratic mean diameter was significantly greater in Thin and Thin + Burn than in Control and Burn treatments. Thinning shifted diameter distributions from uneven- to even-sized. Crown fuel load (CFL) of both pinyon and juniper was significantly lower in Thin and Thin + Burn compared with Control and Burn treatments. Thin, Burn, and Thin + Burn treatments resulted in significantly greater 1-h surface fuel loads compared with the Control. The Thin treatment resulted in significantly greater mean load of the 1000-h fuel class compared with Burn and Control treatments, but did not differ from Thin + Burn. Forest floor Oi (litter) layer was not significantly affected by the treatments but Oe + Oa (duff) depth was significantly less in the Burn treatment compared with Thin and Control. Live shrubs and tree regeneration showed no differences among treatments. We concluded that thinning and thinning followed by prescribed fire were effective approaches for fuels reduction; however, resulting stand structures may be novel and outside the historical range of variability. Prescribed fire alone had minimal effects on structure and fuels reduction. Woody shrubs and tree regeneration in the understory suggested that these treatments may not have long-term deleterious ecological effects.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

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

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

Overstory and shrub effects on natural regeneration processes in native Pinus radiata stands

Native Monterey pine (Pinus radiata D. Don) stands on the Monterey peninsula have been significantly modified by natural and anthropogenic disturbances. The exclusion of fire and the introduction of pitch canker (caused by Fusarium circinatum Nirenberg & O’Donnell) created a need for the examination of these changing ecosystems with respect to regeneration, especially as these stands reach mature ages. We established 210 plots on 35 transects distributed throughout five stands in order to describe the current stand structure and quantify the extent and condition of regeneration. The results indicated that Monterey pine seedling establishment varies throughout the peninsula depending on percent canopy cover, duff and litter depth, and percent shrub cover while seedling growth is influenced by percent shrub cover. Canopy cover was also found to inversely influence shrub cover. Our study highlights the importance of understory removal to increase Monterey pine regeneration and seedling growth as canopy cover decreases, especially in areas where coast live oak (Quercus agrifolia Nee) and poison-oak (Toxicodendron diversilobum E. Greene) are abundant.     

Response of soricid populations to repeated fire and fuel reduction treatments in the southern Appalachian Mountains

Fuel hazards have increased in forests across the United States because of fire exclusion during the 20th century. Treatments used to reduce fuel buildup may affect wildlife, such as shrews, living on the forest floor, especially when treatments are applied repeatedly. From mid-May to mid-August 2006 and 2007, we used drift fences with pitfall traps to capture shrews in western North Carolina in 3 fuel reduction treatment areas [(1) twice-burned (2003 and 2006), (2) mechanical understory cut (2002), and (3) mechanical understory cut (2002) followed by 2 burns (2003 and 2006)] and a control. We captured 77% fewer southeastern shrews (Sorex longirostris) in mechanical + twice-burned treatment areas than in mechanical treatment areas in 2006, but southeastern shrew captures did not differ among treatment areas in 2007. Total shrew captures did not differ among treatment areas in either year. Decreases in leaf litter, duff depth, and canopy cover in mechanical + twice-burned treatment areas may have decreased ground-level moisture, thereby causing short-term declines in southeastern shrew captures. Prescribed fire or mechanical fuel reduction treatments in the southern Appalachian Mountains did not greatly affect shrew populations, though the combination of both treatments may negatively affect some shrew species, at least temporarily.     

Restoring longleaf pine (Pinus palustris Mill.) in loblolly pine (Pinus taeda L.) stands: Effects of restoration treatments on natural loblolly pine regeneration

Historical land use and management practices in the southeastern United States have resulted in the dominance of loblolly pine (Pinus taeda L.) on many upland sites that historically were occupied by longleaf pine (Pinus palustris Mill.). There is currently much interest in restoring high quality longleaf pine habitats to such areas, but managers may also desire the retention of some existing canopy trees to meet current conservation objectives. However, fast-growing natural loblolly pine regeneration may threaten the success of artificially regenerated longleaf pine seedlings. We evaluated the establishment and growth of natural loblolly pine regeneration following different levels of timber harvest using single-tree selection (Control (uncut, residual basal area ∼16 m²/ha), MedBA (residual basal area of ∼9 m²/ha), LowBA (residual basal area of ∼6 m²/ha), and Clearcut (complete canopy removal)) and to different positions within canopy gaps (approximately 2800 m²) created by patch cutting at two ecologically distinct sites within the longleaf pine range: Fort Benning, GA in the Middle Coastal Plain and Camp Lejeune, NC in the Lower Coastal Plain. The density of loblolly pine seedlings was much higher at Camp Lejeune than at Fort Benning at the end of the first growing season after harvesting. Following two growing seasons, there were no significant effects of canopy density or gap position on the density of loblolly pine seedlings at either site, but loblolly pine seedlings were taller on treatments with greater canopy removal. Prescribed fires applied following the second growing season killed 70.6% of loblolly pine seedlings at Fort Benning and 64.3% of seedlings at Camp Lejeune. Loblolly pine seedlings were generally less than 2 m tall, and completeness of the prescribed burns appeared more important for determining seedling survival than seedling size. Silvicultural treatments that include canopy removal, such as patch cutting or clearcuts, will increase loblolly pine seedling growth and shorten the window of opportunity for control with prescribed fire. Therefore, application of prescribed fire every 2-3 years will be critical for control of loblolly pine regeneration during restoration of longleaf pine in existing loblolly pine stands.     

Can prescribed fire be used to maintain fuel treatment effectiveness over time in Black Hills ponderosa pine forests?

We determine the time frame after initial fuel treatment when prescribed fire will be likely to produce high enough mortality rates in ponderosa pine (Pinus ponderosa var. scopulorum Dougl. ex Laws.) regeneration to be successful in maintaining treatment effectiveness in the Black Hills of South Dakota. We measured pine regeneration in disturbed stands and young pine growth rates to estimate the susceptibility of pine regeneration to prescribed fire with time since initial treatment. We also determined surface fuel accumulation rates for stands after prescribed fire to help estimate likely fire behavior in maintenance prescribed fire. Given our estimates of regeneration density and tree size, and likely fire behavior, we then used small pine tree mortality—fire effect relations to estimate the effects of prescribed fire on developing understory pine at specific times since initial treatments.     

Understory vegetation in old-growth and second-growth Tsuga canadensis forests in western Massachusetts

We compared the understory communities (herbs, shrubs, and tree seedlings and saplings) of old-growth and second-growth eastern hemlock forests (Tsuga canadensis) in western Massachusetts, USA. Second-growth hemlock forests originated following clear-cut logging in the late 1800s and were 108–136 years old at the time of sampling. Old-growth hemlock forests contained total ground cover of herbaceous and shrub species that was approximately 4 times greater than in second-growth forests (4.02 ± 0.41%/m² versus 1.06 ± 0.47%/m²) and supported greater overall species richness and diversity. In addition, seedling and sapling densities were greater in old-growth stands compared to second-growth stands and the composition of these layers was positively correlated with overstory species composition (Mantel tests, r > 0.26, P < 0.05) highlighting the strong positive neighborhood effects in these systems. Ordination of study site understory species composition identified a strong gradient in community composition from second-growth to old-growth stands. Vector overlays of environmental and forest structural variables indicated that these gradients were related to differences in overstory tree density, nitrogen availability, and coarse woody debris characteristics among hemlock stands. These relationships suggest that differences in resource availability (e.g., light, moisture, and nutrients) and microhabitat heterogeneity between old-growth and second-growth stands were likely driving these compositional patterns. Interestingly, several common forest understory plants, including Aralia nudicaulis, Dryopteris intermedia, and Viburnum alnifolium, were significant indicator species for old-growth hemlock stands, highlighting the lasting legacy of past land use on the reestablishment and growth of these common species within second-growth areas. The return of old-growth understory conditions to these second-growth areas will largely be dependent on disturbance and self-thinning mediated changes in overstory structure, resource availability, and microhabitat heterogeneity.     

Spatial pattern in herb diversity and abundance of second growth mixed deciduous-evergreen forest of southern New England,USA

This study was designed to answer questions about the patterns of understory diversity in managed forests of southern New England, and the factors that appear associated with those patterns. At the landscape-level, we used plot data to answer questions regarding the spatial distribution of forest understory plant species. Data from a combination of fixed area (understory vegetation) and variable radius (overstory trees) plot methods are combined with site variables for the analysis. Univariate and multivariate statistical methods are used to test for understory diversity relationships with overstory cover types and topography separately, and in combination. Analyses also test for relationships between specific understory species and cover types. In general the understory flora is dominated by four common clonal species that occur across the range of forest cover types: wild sarsaparilla (Aralia nudicaulis L.), Canada mayflower (Maianthemum candense Desf.), star flower (Trientalis borealis Raf.), and partridgeberry (Mitchella repens L.). Results also show that over story composition and structure can be used to assess understory species richness. Species richness follows a general trend among cover types of: hardwood ≥ regenerating forest, hardwood–pine, and pine ≥ mixed ≥ hardwood–hemlock > hemlock. Eastern hemlock (Tsuga canadensis L. Carriere) and mountain laurel (Kalmia latifolia L.) (which decreased in dominance from ridge to valley) both showed negative trends with understory species richness. Topographic position also appears associated with understory floristic patterns (particularly for the hardwood cover type), both in terms of species richness and compositional diversity which both increased from ridge, to midslope, to valley. However, overstory composition (covertype) appears to have a higher order influence on vegetation and mediates the role of topography. The results from this study provide foresters with a better understanding for maintaining floristic diversity and composition of the understory in managed forests.     

Tree regeneration and future stand development after bark beetle infestation and harvesting in Colorado lodgepole pine stands

In the southern Rocky Mountains, current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks and associated harvesting have set millions of hectares of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) forest onto new stand development trajectories. Information about immediate, post-disturbance tree regeneration will provide insight on dynamics of future stand composition and structure. We compared tree regeneration in eight paired harvested and untreated lodgepole pine stands in the Fraser Experimental Forest that experienced more than 70% overstory mortality due to beetles. New seedlings colonized both harvested and untreated stands in the first years after the beetle outbreak. In harvested areas the density of new seedlings, predominantly lodgepole pine and aspen, was four times higher than in untreated stands. Annual height growth of pine and fir advance regeneration (e.g., trees established prior to the onset of the outbreak) has doubled following overstory mortality in untreated stands. Growth simulations based on our regeneration data suggest that stand basal area and stem density will return to pre-beetle levels in untreated and harvested stands within 80-105 years. Furthermore, lodgepole pine will remain the dominant species in harvested stands over the next century, but subalpine fir will become the most abundant species in untreated areas. Owing to terrain, economic and administrative limitations, active management will treat a small fraction (<15%) of the forests killed by pine beetle. Our findings suggest that the long-term consequences of the outbreak will be most dramatic in untreated forests where the shift in tree species composition will influence timber and water production, wildfire behavior, wildlife habitat and other forest attributes.     

Small-scale variation of vegetation in a mixed forest understorey is partly controlled by the effect of overstory composition on litter accumulation

We investigated how richness and composition of vascular plant species in the understory of a mixed hardwood forest stand varied with respect to the abundance and composition of the overstory. The stand is in central Spain and represents the southernmost range of distribution of several tree and herbaceous species in Europe. Understory species were identified in 46 quadrats (0.25 m²) where variables litter depth and light availability were measured. In addition, we estimated tree density, basal area, and percent basal area by tree species within 6-m-radius areas around each plot. Species richness and composition were studied using path analysis and scale-dependent geostatistical methods, respectively. We found that the relative abundance of certain trees species in the overstory was more important than total overstory abundance in explaining understory species richness. Richness decreased as soil litter depth increased, and soil litter increased as the relative proportion of Fagus sylvatica in the overstory increased, which accounted for a negative, indirect effect of Fagus sylvatica on richness. Regarding understory species composition, we found that some species distributed preferentially below certain tree species. For example, Melica uniflora was most frequent below Fagus sylvatica and Quercus petraea while the increasing proportion of Q. pyrenaica in the overstory favored the presence of Cruciata glabra, Arenaria montana, Prunus avium, Conopodium bourgaei, Holcus mollis, Stellaria media and Galium aparine in the understory. Overall, these results emphasize the importance of individual tree species in controlling the assemblage and richness of understory species in mixed stands. We conclude that soil litter accumulation is one way through which overstory composition shapes the understory community.     

Decades-old silvicultural treatments influence surface wildfire severity and post-fire nitrogen availability in a ponderosa pine forest

Wildfire severity and subsequent ecological effects may be influenced by prior land management, via modification of forest structure and lingering changes in fuels. In 2002, the Hayman wildfire burned as a low to moderate-severity surface fire through a 21-year pine regeneration experiment with two overstory harvest cuttings (shelterwood, seed-tree) and two site preparations (scarified, unscarified) that had been applied in a mature ponderosa pine forest in the montane zone of the Colorado Front Range in 1981. We used this event to examine how pre-fire fine fuels, surface-level burn severity and post-fire soil nitrogen-availability varied with pre-fire silvicultural treatments. Prior to the wildfire, litter cover was higher under both shelterwood and unscarified treatments than seed-tree and scarified treatments. Immediately after the fire in 2002, we assessed burn severity under 346 mature trees, around 502 planted saplings, and in 448 4 m² microplots nested within the original experimental treatments. In one-fourth of the microplots, we measured resin-bound soil nitrate and ammonium accumulated over the second and third post-fire growing season. Microplots burned less severely than bases of trees and saplings with only 6.8% of microplot area burned down to mineral soil as compared to >28% of tree and sapling bases. Sapling burn severity was highest in unscarified treatments but did not differ by overstory harvest. Microplot burn severity was higher under the densest overstory (shelterwood) and in unscarified treatments and was positively related to pre-fire litter/duff cover and negatively associated with pre-fire total plant cover, grass cover and distance to tree. In both years, resin-bound nitrate and ammonium (NH₄⁺-N) increased weakly with burn severity and NH₄⁺-N availability was higher in unscarified than scarified plots. The lasting effects of soil scarification and overstory harvest regime on modern patterns of surface burn severity after two decades underscores the importance of historic landuse and silviculture on fire behavior and ecological response. Unraveling causes of these patterns in burn severity may lead to more sustainable fire and forest management in ponderosa pine ecosystems.     

The burning characteristics of southeastern oaks: Discriminating fire facilitators from fire impeders

In southeastern pine–oak ecosystems, ecological restoration targets oaks for removal by chemical, mechanical, burning, or combinations of treatments. Managers often pursue oak removal indiscriminately despite the poorly understood historical structure, cover, and ecological function within these ecosystems. Restoration treatments often cite the impediment that oak litter represents to prescribed fire spread and effectiveness. We evaluated the burning characteristics of eight southeastern Quercus spp. by burning collected litter under controlled conditions in a combustion chamber. Replicated burns consisted of 15 g of litter on a 35 cm × 35 cm grid of xylene-soaked cotton strings. Burning characteristics measured included maximum flame height (cm), flaming duration (s), smoldering duration (s), residual ash, and mass loss rate (g s⁻¹). We compared all 8 oaks using ANCOVA, with litterbed depth as a covariate. The oaks differed for all burning characteristics measured (P < 0.001). Rank comparisons placed Quercus stellata and Quercus laevis as the species with greatest fire intensity, sustainability, and consumability, equivalent in many measures to longleaf pine and other fire resisters. Quercus virginiana and Quercus hemisphaerica burned with the least intensity, sustainability, and consumability, burning similarly to sand pine and other fire evaders. These results show that oaks common to southeastern United States ecosystems have litter properties, similar to pines, which vary in their ability to sustain fire. Understanding the pyric properties of oak species also suggests that managers prioritize removal of species that hinder prescribed fire effectiveness for restoration of southeastern USA pine–oak ecosystems.