The effects of prescribed burning and canopy openness on establishment of two non-native plant species in a deciduous forest,southeast Ohio,USA

Fire has often been shown to promote invasion by non-native plant species, but few studies have examined the process in temperate-zone deciduous forests. To examine the potential of prescribed fire to facilitate invasions in the Central Hardwoods ecosystem, we experimentally burned small plots and simulated aspects of fire at a forested site in southeastern Ohio, USA. Treatments included high and low burn intensity, lime addition, and litter removal to test hypotheses of population limitation by fire intensity, fire-caused nutrient release, and removal of leaf litter, respectively. Treatments were arranged in a randomized block design in two landscape positions (dry upland, moist lowland) and two canopy conditions (gap, no gap). The experimental sites were not significantly different from randomly chosen forest sites in any of 12 environmental variables. Seeds of two problematic non-native species (Microstegium vimineum and Rosa multiflora) were sown into plots following treatment to test the possibility of seed limitation. We recorded germination and height growth at three dates 1, 4, and 14 months following burning. Germination was promoted by litter removal and high- and low-intensity fire treatments in M. vimineum, and by high-intensity fire in R. multiflora. Seedling growth of both species was greatest following high-intensity fire under canopy gaps. Germination in the second year showed treatment effects similar to the first year indicating persistence of fire effects. Both species showed stronger recruitment in valleys and in canopy gaps, reflecting an interaction of fire and landscape position. We infer that prescribed burning and canopy-opening management practices have the potential to facilitate invasion of the study area by creating conditions promoting establishment and growth of at least two non-native species. The absence of these species in previous studies appears to be due to a lack of propagules rather than the unsuitability of forest sites for germination or growth.     

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.     

The role of disturbance severity and canopy closure on standing crop of understory plant species in ponderosa pine stands in northern Arizona,USA

Concerns about the long-term sustainability of overstocked dry conifer forests in western North America have provided impetus for treatments designed to enhance their productivity and native biodiversity. Dense forests are increasingly prone to large stand-replacing fires; yet, thinning and burning treatments, especially combined with other disturbances such as drought and grazing, may enhance populations of colonizing species, including a number of non-native species. Our study quantifies plant standing crop of major herbaceous species across contrasting stand structural types representing a range in disturbance severity in northern Arizona. The least disturbed unmanaged ponderosa pine stands had no non-native species, while non-native grasses constituted 7–11% of the understory plant standing crop in thinned and burned stands. Severely disturbed wildfire stands had a higher proportion of colonizing native species as well as non-native species than other structural types, and areas protected from grazing produced greater standing crop of native forbs compared to grazed unmanaged stands. Standing crop of understory plants in low basal area thinned and burned plots was similar to levels on wildfire plots, but was comprised of fewer non-native graminoids and native colonizing plants. Our results also indicate that size of canopy openings had a stronger influence on standing crop in low basal area plots, whereas tree density more strongly constrained understory plant standing crop in dense stands. These results imply that treatments resulting in clumped tree distribution and basal areas <10 m² ha⁻¹ will be more successful in restoring native understory plant biomass in dense stands. Multiple types and severity of disturbances, such as thinning, burning, grazing, and drought over short periods of time can create greater abundance of colonizing species. Spreading thinning and burning treatments over time may reduce the potential for non-native species colonization compared to immediately burning thinned stands.     

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.     

Fire-vegetation interplay in a mosaic structure of Quercus suber woodlands and Mediterranean maquis under recurrent fires

We examined the effects of fire recurrence on a mosaic structure of Quercus suber and Erica-Cistus shrubland communities of Southeastern France in order to improve the understanding of fire-vegetation interplay. Plant communities that were similar in 1959 (woodlands on shrublands called maquis on acidic soils) were compared along a gradient of fire recurrence, from 0 to 4 fires, with different time intervals between fires. The results showed that understory cover increased roughly with fire recurrence, whereas tree height, cover, density, stand basal area and litter depth decreased. Different fire recurrences along the past decades led to different vegetation types. High fire recurrence corresponded to maquis and sparse cork-oak woodlands while pure oak woodlands established in the absence of fire during the same period. In all, the diversity of tree diameter decreased with fire recurrence. High fire recurrence (3 or 4 fires in 50 years) led to a simple vertical structure of vegetation with a mono-layered shrub cover and few Quercus suber trees. In contrast, spatial connections between plants were maximal at longer time intervals, leading to a multi-layered vegetation. We finally discuss the potential implications of past fires on the behavior of future fires in the perspective of a sustainable management of these Mediterranean ecosystems.     

Long-term effects of single prescribed fires on hardwood regeneration in oak shelterwood stands

One of the arguments against using prescribed fire to regenerate oak (Quercus spp.) forests is that the improvement in species composition of the hardwood regeneration pool is temporary and multiple burns are necessary to achieve and maintain oak dominance. To explore this concern, I re-inventoried a prescribed fire study conducted in the mid-1990s to determine the longevity of the effects of a single prescribed fire on hardwood regeneration. The initial study was conducted in three oak shelterwood stands in central Virginia, USA. In 1994, each stand was divided into four treatments (spring, summer, and winter burns and a control) and the hardwood regeneration was inventoried before the fires. During the burns, fire intensity was measured and categorized in each regeneration sampling plot. Second-year postfire data showed marked differences in species mortality rates, depending on season-of-burn and fire intensity: oak and hickory (Carya spp.) regeneration dominated areas burned by medium- to high-intensity fire during the spring and summer while yellow-poplar (Liriodendron tulipifera) and red maple (Acer rubrum) seedlings dominated unburned areas and all areas treated with low-intensity fire regardless of season-of-burn. The treatments were re-inventoried in 2006 and 2007 to determine whether these fire effects were still present. The new data show that the species distributions by season-of-burn and fire intensity found in 1996 still existed 11 years after the treatments. The fact that fire effects in oak shelterwood stands can last at least a decade has important management implications for resource professionals interested in sustaining oak forests in the eastern United States.     

Prescribed burning and understory composition in a temperate deciduous forest,Ohio, USA

Since the advent of widespread suppression in the mid-20th century, fire has been relatively rare in deciduous forests of the eastern United States. However, widespread prescribed burning has recently been proposed as a management tool to favor oak (Quercus spp.) regeneration. To examine the potential effects of fire introduction on the understory community, we experimentally burned small plots and simulated aspects of fire at a forested site in southeastern Ohio. Treatments included two burn intensities, litter removal, increased soil pH, and a control. Treatments were arranged in a randomized block design in two landscape positions (dry upland and moist lowland) and two canopy conditions (gap, no gap). Post-fire vegetation was identified to species, and stems were counted 1, 3, and 14 months after burning. Community composition was more strongly affected by fire in upland plots than in lowlands, but was not affected by canopy openness. Both cool and hot burns reduced post-fire seedling emergence of Acer rubrum, a common overstory tree. Hot burns facilitated germination of Vitis spp., Rhus glabra, and Phytolacca americana, species common in disturbed habitats, and increased graminoid abundance. Cool burns and litter removal facilitated germination of Erechtites hieracifolia and Liriodendron tulipifera suggesting that litter removal is the mechanism by which fire favors colonization. These results suggest that fire applied frequently in the Central Hardwoods Region would cause compositional shifts to graminoids and disturbance-adapted forbs by increasing germination from the seed bank. Fire did not favor species with dormant underground buds, as studies in other ecosystems would suggest. Vegetational responses were noticeably weaker in the second year after burning, indicating that a single fire has only a short-term effect.     

Ecological consequences of an exotic fungal disease in eastern U.S. hardwood forests

Exotic pests and pathogens can cause extensive mortality of native species resulting in cascading effects within an ecosystem. As ecosystems lose species to exotic enemies, ecosystem function may be disrupted if the ecological roles are not filled by the remaining species. To illustrate this concept, this paper examines the impacts of an exotic fungus (Discula dectructiva) on flowering dogwood (Cornus florida), historically a common understory tree species in eastern U.S. hardwood forests. Recent studies indicate that dogwood plays an important role in the health and ecological integrity of forest ecosystems throughout the eastern U.S. by increasing the availability of calcium in the biota-rich surface horizons of forest soils. However, Discula destructiva causes a disease, dogwood anthracnose, which can rapidly kill dogwood trees. This paper also illustrates how past fire has increased dogwood density and improved tree health in areas infected with anthracnose, suggesting that prescribed fire may offer a tool for land managers to maintain dogwood as a component in eastern U.S. hardwood forests by shifting the “ideal” disturbance regime of this previously fire-intolerant species.     

Can pine forest restoration promote a diverse and abundant understory and simultaneously resist nonnative invasion?

An important goal of forest restoration is to increase native plant diversity and abundance. Thinning and burning treatments are a common method of reducing fire risk while simultaneously promoting understory production in ponderosa pine (Pinus ponderosa) forests. In this study we examine the magnitude and direction of understory plant community recovery after thinning and burning restoration treatments in a ponderosa pine forest. Our objective was to determine if the post-treatment community was a diverse, abundant, and persistent assemblage of native species or if ecological restoration treatments resulted in nonnative species invasion. This project was initiated at the Grand Canyon-Parashant National Monument, Arizona, USA in 1997. We established four replicated blocks that spanned a gradient of soil types. Each block contained a control and a treated unit. Treated units were thinned to emulate pre-1870 forest stand conditions and prescribed-burned to reintroduce fire to a system that has not burned since ∼1870. We measured plant cover using the point-line intercept method and recorded species richness and composition on 0.05 ha belt transects. We examined the magnitude of treatment responses using Cohen's d effect size analysis. Changes in community composition were analyzed using nonmetric multidimensional scaling (NMS). Native plant species cover and richness increased in the thinned and burned areas compared to the controls. By the last year of the study, annual species comprised nearly 60% of the understory cover in the treatment units. Cheatgrass (Bromus tectorum), a nonnative annual grass, spread into large areas of the treated units and became the dominant understory species on the study site. The ecological restoration treatments did promote a more diverse and abundant understory community in ponderosa pine forests. The disturbances generated by such treatments also promoted an invasion by an undesirable nonnative species. Our results demonstrate the need to minimize disturbances generated by restoration treatments and argue for the need to proactively facilitate the recovery of native species after treatment.     

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.     

Burning and mowing as habitat management for capercaillie Tetrao urogallus: An experimental test

In Scots pine Pinus sylvestris forests, the important ecological effects of natural fires could be emulated using prescribed fire. Species that may benefit from fire effects include capercaillie Tetrao urogallus, a large forest grouse. A key component of forest habitats for capercaillie is the ericaceous shrub, bilberry Vaccinium myrtillus, which is eaten by capercaillie, and supports abundant arthropods, taken by young chicks. We carried out an experiment testing whether prescribed burning would be a valuable technique for capercaillie habitat management. The study took place at Abernethy Forest, the largest ancient native pinewood in Britain, and a key capercaillie site, holding c 8-20% of the British population. Prescribed fire in woodland is highly novel in Britain. We therefore also tested mowing, which might replicate some fire effects more cheaply and safely. Twenty-five experimental blocks were established within open pine stands with ground vegetation including bilberry, but dominated by heather Calluna vulgaris. Each block held three 700 m² plots, randomly assigned to control, mow and burn. Vegetation, arthropods and capercaillie dung were monitored over a 7-year period, including 1 year prior to treatment. Mean bilberry cover, initially around 12%, increased in mown and burnt areas, but there were also increases in controls, following unusual natural die-back of heather. By the sixth season after treatment, bilberry cover was significantly higher in burnt and mown areas than controls, averaging 27% (95% confidence intervals 24-30), compared to 20% (19-21) in controls. Biomass of spiders, an important dietary group for capercaillie chicks, as measured by pitfall trapping, was significantly higher in burnt and mown plots than controls, by about 56% (38-76). However, biomass of caterpillars, often considered a more important dietary group, did not show clear differences between treatments. An alternative analysis was used to ‘statistically remove’ natural heather die-back; this enhanced the treatment differences in bilberry cover and spider biomass. Capercaillie dung counts suggested that burnt, and especially mown areas, had more summer capercaillie usage than controls. Capercaillie conservation at sites similar to Abernethy is likely to benefit from either prescribed fire or mowing, because these techniques increase bilberry and spider abundance. This study illustrates the value of collaboration between researchers and land-managers, in developing and testing novel management techniques. We support the idea that ‘dominance reduction’, delivered through managed disturbance, offers a general principle to guide land-managers wishing to maintain biodiversity, particularly where key species, like capercaillie, are strongly associated with sub-dominant plant species like bilberry.     

Post-fire germination: The effect of smoke on seeds of selected species from the central Mediterranean basin

In regions with a Mediterranean-type climate wildfires are a frequent occurrence: in such environments fire tolerant/favoured species are frequently encountered. In the Mediterranean basin, many species of fire prone habitats are resprouters while others are known to germinate after fire. Fire causes an enhancement of seed germination in many species from fire prone habitats in the other regions with a Mediterranean-type vegetation such as Western Australia, California and South Africa. Seeds of a number of these species are stimulated to germinate by the smoke generated from burning of plant material in either an aerosol or aqueous form. However, for species from the Mediterranean basin the role of smoke in germination is poorly known, despite the fact that in the field many species seems to be encouraged to germinate after fire. We examined the germination of 10 species native to the Mediterranean basin that were treated with aerosol smoke. Some species were from fire prone habitats while others were not. In relation to the controls, increased germination occurred in three of the species (e.g. Cistus incanus), three had more rapid germination but no total increase (e.g. Rhamnus alaternus), two showed reduced germination (e.g. Asphodelus ramosus) and two exhibited no difference in germination (e.g. Clematis flammula). There was additionally no consistent pattern of germination behaviour depending on the habitat from which the species came. Comparison is made between the results of this study and those of other studies on seed germination response to heat and smoke in other areas of Mediterranean-type climate. An understanding of the importance of fire in relation to other disturbances in the vegetation dynamics in the Mediterranean basin needs to be clarified by further detailed studies of the effect of heat and smoke products on seed germination of Mediterranean species. Outcomes of further research, also on a broader range of species, would have important impacts also for conservation, environment management, horticulture and ecosystem restoration.     

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.     

Bird communities following high-severity fire: Response to single and repeat fires in a mixed-evergreen forest,Oregon, USA

Fire is a widespread natural disturbance agent in most conifer-dominated forests. In light of climate change and the effects of fire exclusion, single and repeated high-severity (stand-replacement) fires have become prominent land management issues. We studied bird communities using point counting in the Klamath-Siskiyou ecoregion of Oregon, USA at various points in time after one or two high-severity fires. Time points included 2 and 3 years after a single fire, 17 and 18 years after a single fire, 2 and 3 years after a repeat fire (15 year interval between fires), and >100 years since stand-replacement fire (mature/old-growth forest). Avian species richness did not differ significantly among habitats. Bird density was highest 17 and 18 years after fire, lowest 2 years after fire, and intermediate in repeat burns and unburned forest. Bird community composition varied significantly with habitat type (A = 0.24, P < 0.0001) with two distinct gradients in species composition relating to tree structure (live to dead) and shrub stature. Using indicator species analysis, repeat burns were characterized by shrub-nesting and ground-foraging bird species while unburned mature forests were characterized by conifer-nesting and foliage-gleaning species. Bird density was not related to snag basal area but was positively related to shrub height. Contrary to expectations, repeated high-severity fire did not reduce species richness, and bird densities were greater in repeat burns than in once-burned habitats. Broad-leaved hardwoods and shrubs appear to play a major role in structuring avian communities in the Klamath-Siskiyou region. In light of these results, extended periods of early seral broadleaf dominance and short-interval high-severity fires may be important to the conservation of avian biodiversity.     

Effects of light/darkness, thermal shocks and inhibitory components on germination of Pinus canariensis, Pinus halepensis and Pinus pinea

Knowledge of the germination characteristics of Pinus species can help in the understanding, prediction and management of the regeneration of pine forests. In the Canary Islands, several exotic pines (Pinus halepensis L. and Pinus pinea L.) were planted with Pinus canariensis Chr. Sm. Ex DC, the only native pine species, and there is now an interest in controlling these exotic species to restore the original forest. The main objective of the present study is to determine the germination response of P. canariensis, P. halepensis and P. pinea to different light regimes (darkness and light/darkness), thermal shocks and the presence of inhibitory substances from the leaves and litter of P. canariensis. P. halepensis seeds showed the highest viability and germination rate. Darkness accelerated P. halepensis germination, while exudates accelerated P. canariensis germination. Only treatments of 200 °C for 5 min and 300 °C for 5 min significantly decreased the germination of all three pine species. In the absence of strong differences in germination among species after treatments, the key for fire management or prescribed burning in this case may be the sprouting ability of P. canariensis. A possible management strategy to control the two obligate seeder exotic species could be to use one intense prescribed fire followed by a low-intensity prescribed fire after seed bank germination in the field, so in this way, only P. canariensis could resprout after fire and it would eliminate only the exotic pines in a mixed stand.     

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.     

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.     

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.     

Toward reference conditions: wildfire effects on flora in an old-growth ponderosa pine forest

Remote ponderosa pine (Pinus ponderosa) forests on the North Rim of Grand Canyon National Park, Arizona, USA provide valuable examples of reference conditions due to their relatively uninterrupted fire regimes, limited grazing history, and protection from logging. Wildfire is an important disturbance agent in upland forests of the Interior West, yet repeated measurements taken before and after lightning-ignited fires are rare. In 1999, a low-severity Wildland Fire Use fire burned 156 ha on Fire Point, a peninsula dominated by old-growth ponderosa pines, which had not burned for at least 76 years. We measured understory plant community and forest floor characteristics in 1998 (1 year before the fire) and 2001 (2 years after the fire) at this site and at nearby reference sites that did not burn in 1999 but have had continuing fire regimes throughout the past century. After the wildfire, the plant community at Fire Point shifted toward higher compositional similarity with the reference sites. Analysis of functional group composition indicated that this change was due primarily to an increase in annual and biennial forbs. Gayophytum diffusum, Polygonum douglasii, Chenopodium spp., Solidago spp., Elymus elymoides, Calochortus nuttallii, Hesperostipa comata, and Lotus spp. were indicative of forests influenced by recent fires. Species richness, plant cover, plant layer density and plant diversity were significantly lower at Fire Point than at the reference sites, possibly due to long-term fire exclusion, but the fire did not increase the rate of change in these variables after 2 years. Few exotic species were present at any site. Forest floor depths at Fire Point were reduced to depths similar to the reference sites, primarily due to consumption of the duff layer. There was a significant inverse relationship between the ratio of duff:litter and species richness. Compared to fire-excluded forests, old-growth ponderosa pine forests influenced by low-intensity surface fires generally have greater plant species richness (especially annual forbs) and lighter fuel loads. This study supports the continued application of the Wildland Fire Use strategy in old-growth montane forests to maintain and improve forest health by altering understory species composition and reducing fuel loads.