Litter flammability in oak woodlands and shrublands of southeastern France

Characterizing the flammability of litter fuels is of major importance for assessing wildland fire ignition hazard. Here we compared the flammability of litter within a mosaic of Quercus suber (cork oak) woodlands and shrublands in a Mediterranean fire-prone area (Maures massif, southeastern France) to test whether the characteristics and the flammability of litter vary with the vegetation types. We tested experimentally the ignitability, the sustainability, the combustibility and the consumability of undisturbed (=non-reconstructed) litter samples with a point-source mode of ignition. Although the frequency of ignition was similar between all the vegetation types, we distinguished four groups having litter of specific composition and flammability: low and sparse shrublands dominated by Cistus species, medium shrublands with cork oak, high Erica shrublands with sparse cork oak woodlands, and mixed mature oak woodlands with Q. suber, Q. ilex and Q. pubescens. As these vegetation types corresponded to a specific range of past fire recurrence, we also tested the effect of the number of fires and the time since the last fire on litter flammability. Litters of plots recurrently burned had low ability to propagate flames and low flame sustainability. We discuss how the recent fire history can modify vegetation and litter flammability, and thus the fire ignition hazard.     

Establishment and growth of oak (Quercus alba, Quercus prinus) seedlings in burned and fire-excluded upland forests on the Cumberland Plateau

Recurrent problems with regeneration of oaks (Quercus spp.) have been documented across a wide range of ecosystems. In oak-dominated forests of the central and Appalachian hardwood regions of the United States, a lack of competitive oak regeneration has been tied, in part, to fire suppression in these landscapes, and managers throughout the region are using prescribed fire to address this concern. To examine fire effects on oak regeneration, researchers have generally relied on inventories or population studies of existing seedlings. These studies are valuable but do not permit examination of the role of fire in enhancing the establishment and growth of new oak seedlings stemming from oak mast events. In this study, white (Quercus alba) and chestnut oak (Quercus prinus) acorn mast crops serendipitously occurred in year three (fall 2005) of a landscape-scale prescribed fire experiment. We examined establishment, survival, height and diameter of new seedlings on sites on the Cumberland Plateau in eastern Kentucky. Treatments were fire exclusion, a single prescribed fire (1x-burn; 2003), and repeated prescribed fire (3x-burn; 2003, 2004, and after acorn drop in 2006), all conducted in late spring. Initial densities of newly established chestnut and white oak seedlings were statistically similar across treatments (P = 0.42), despite fires on the 3x-burn site having occurred after acorns were on the ground. Oak seedling density was significantly predicted by oak basal area on all sites (R² = 0.12–0.46), except for chestnut oak on fire-excluded sites (R² = 0.04). Litter depth was less on 3x-burn sites compared to 1x-burn and fire-excluded sites, whereas canopy openness was greater on both burn treatments compared to fire-excluded sites. Seedling mortality was generally higher on fire-excluded sites compared to burn sites, especially for white oak. Oak seedling mortality in the first two growing seasons was significantly predicted by initial litter depth and open sky, with greater litter depth and lower percent open sky leading to higher mortality. In the third growing season none of the measured variables predicted chestnut oak seedling survival; for white oak, percent open sky remained a significant predictor of mortality. Initially, seedlings on the fire-excluded sites had similar height but smaller diameter; after three growing seasons there were few differences in seedling height or diameter among treatments. Our findings suggest a potential role for prescribed fire in establishing forest floor and light conditions that may enhance the success of new oak germinants, although different responses among species may suggest the need to target management for individual oak species.     

Fire-driven oak regeneration in French Mediterranean ecosystems

Regeneration by seeds for cork oak (Quercus suber) and companion oaks (holm oak Quercus ilex and downy oak Quercus pubescens) is likely to be poor in the fire-prone Maures massif (southern France) but the causes are poorly known. Our objective was to assess the effective recruitment for these three oak species and their temporal pattern of recruitment, in order to determine the main limitation factors and the regeneration window of each species. We studied oak recruits (height <3 m) in naturally regenerated populations according to a gradient of fire recurrence and in five main vegetation types including shrublands and mixed mature woodlands. Fire recurrence was the main explanatory factor of oak recruitment, either directly or through vegetation type and microsite characteristics. The results indicate nil to low recruitment for holm oak and downy oak in shrublands, especially those recurrently burned and dominated by Cistus species. Cork oak recruited better than the other oaks in medium and high shrublands dominated by Erica arborea. In contrast, recruitment was high for holm and downy oak in mixed oak stands and mixed pine-oak stands that have not burned for decades. Microsite conditions such as coverage by litter and shrubs influenced oak recruitment, whereas landscape configuration and stand basal area had no influence. Our results suggest that strategic shrub-clearing, oak planting and protection of mixed oak woodlands as seed sources would help maintaining oak populations in the woodland–shrubland mosaic.     

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.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

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.     

Oak regeneration using the shelterwood-burn technique: management options and implications for songbird conservation in the southeastern United States

Shelterwood silviculture is commonly used to regenerate oaks in upland stands. However, competition from other species such as tulip-poplar (Liriodendron tulipifera) may deter oak regeneration when these traditional shelterwood techniques are used. The shelterwood-burn technique is a relatively new tool for regenerating oak-dominated stands on some upland sites while simultaneously minimizing undesirable hardwood intrusion with prescribed fire. Once successful oak regeneration has been achieved, three options are available which will result in different vegetative structure and composition within a stand and subsequently different habitats for songbirds. These options are: complete or partial canopy retention, post-harvest prescribed burning and complete canopy removal. Canopy retention, burning and removal treatments will create, respectively, two-age stands that are likely to harbor a diverse mixture of mature forest and early successional species; park-like woodlands with open woodland species; or early-successional habitats with shrubland species. We suggest that shelterwood-burn systems and the management options associated with them offer viable alternatives for managing both songbird and timber resources where oak-dominated stands are the desired goal in upland southeastern sites.     

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.     

Effects of vegetation type and fire regime on flammability of undisturbed litter in Southeastern France

To assess the effect of vegetation types and of fire regime on the flammability of dead fuels, samples of litter were collected undisturbed (i.e. keeping the structure of litter layers) in 29 study sites spread over the limestone-derived soils of Provence. The sampling plan comprised the most representative ecosystems of the study area: pure Pinus halepensis stands, mixed pine-oak stands and shrublands. Three classes of litter depth were studied (low, medium and high) to account for the variability existing in the field. Sampling also included the number of fires in each site since 1960 (no fire, 1 or 2 fires and ≥3 fires) and the time interval since the last fire (≤15 years, 15-45 years and >45 years). Flammability experiments were carried out in laboratory using a glowing firebrand and a 10 km h⁻¹ wind. The main variables recorded were: ignition frequency, time-to-ignition, flaming duration, flame rate of spread, flame propagation, mean flame temperature, maximum flame height and rate of consumption.Results showed that the ignition frequency was higher in mixed stands than in pure pine stands whereas the time-to ignition, flaming duration and rate of consumption were the highest in pure pine stands. The maximum flame height and the flame propagation decreased with the increase of the number of fires and the time-to-ignition was the highest when the interval since the last fire was the shortest. Increases in litter depth resulted in increased mean flame temperature, maximum flame height and flame propagation. These results can be explained, in part, by the proportions of the different litter components.     

Oak decline in the Boston Mountains,Arkansas, USA: Spatial and temporal patterns under two fire regimes

A spatially explicit forest succession and disturbance model is used to delineate the extent and dispersion of oak decline under two fire regimes over a 150-year period. The objectives of this study are to delineate potential current and future oak decline areas using species composition and age structure data in combination with ecological land types, and to investigate how relatively frequent simulated fires and fire suppression affect the dynamics of oak decline. We parameterized LANDIS, a spatially explicit forest succession and disturbance model, for areas in the Boston Mountains of Arkansas, USA. Land type distribution and initial species/age class were parameterized into LANDIS using existing forest data. Tree species were parameterized as five functional groups including white oak (Quercus alba L., Quercus stellata Wangenh., Quercus muehlenbergii Engelm.), red oak (Qurecus rubra L., Quercus marilandica Muenchh., Quercus falcata Michx., Quercus coccinea Muenchh.), black oak (Quercus velutina Lam.), shortleaf pine (Pinus echinata Mill), and maple (Acer rubrum L., Acer saccharum Marsh.) groups. Two fire regimes were also parameterized: current fire regime with a fire return interval of 300 years and a historic fire regime with an overall average fire return interval of 50 years. The 150-year simulation suggests that white oak and shortleaf pine abundance would increase under the historic fire regime and that the red oak group abundance increases under the current fire regime. The black oak group also shows a strong increasing trend under the current fire regime, and only the maple group remains relatively unchanged under both scenarios. At present, 45% of the sites in the study area are classified as potential oak decline sites (sites where red and black oak are >70 years old). After 150 simulation years, 30% of the sites are classified as potential oak decline sites under the current fire regime whereas 20% of the sites are potential oak decline sites under the historic fire regime. This analysis delineates potential oak decline sites and establishes risk ratings for these areas. This is a further step toward precision management and planning.     

Gradient analysis of an eastern sand savanna's woody vegetation,and its long-term responses to restored fire processes

Little information is available comparing historic and modern sand savannas, and how remnants respond to restored fire. We compared short- and long-term effects of restored fire on the Tefft Savanna, a 197 ha eastern sand savanna in northwest Indiana that had undergone three decades of fire protection. U.S. Public Land Survey data from Tefft in 1833 indicate black and white oak barrens, and pin oak savanna, with trees averaging 50 stems/ha and 4 m²/ha basal area. We used ordination and a digital elevation model to assess topographic distribution of tree species in 1986. In 1986, we also compared initial effects of high- and low-intensity dormant season fire on woody vegetation among nine blocks containing black oak, white oak, and pin oak stands. Twenty years later, we compared the same blocks, all of which had been burned three times per decade with low-intensity fires. In 1986, black oak, white oak and pin oak occurred across a gradient of decreasing elevation and slope. At that time, unburned black oak and white oak stands averaged >400 stems/ha and about 10 m²/ha basal area, and their smaller size classes contained non-oak woody vegetation that apparently had invaded with fire exclusion. After initial burns, black oak and white oak stands receiving high-intensity fire averaged <200 stems/ha and had significantly lower oak canopy cover and basal area than unburned stands. Stands receiving low-intensity fire had intermediate oak canopy cover, with basal area similar to unburned stands. Pin oak stands were more fire-resistant, apparently because spring flooding often reduced fire effects. Density, cover and basal area of non-oak tree species were much lower than oaks, and were not reduced by initial burning. Repeated low-intensity burning over 20 years tended to maintain structure caused by initial fires. However, it reduced lower size class stem densities, promoted post-fire sprouting into the shrub layer, and allowed oak basal area to increase in larger size classes. Time since fire regulated shrub layer structure on a 4-year cycle. Density and cover of trees and shrubs returned to pre-burn conditions by the second and fourth growing seasons after fire, respectively, with non-oak tree species exceeding pre-burn cover and density by the fourth season. These results suggest that high-intensity fire is more important than repeated low-intensity burning in structuring and restoring eastern sand savanna, and that non-oak tree species, once established, may be resistant to low-intensity fire.     

Does tree species composition control soil organic carbon pools in Mediterranean mountain forests?

We compared soil organic carbon (SOC) stocks and stability under two widely distributed tree species in the Mediterranean region: Scots pine (Pinus sylvestris L.) and Pyrenean oak (Quercus pyrenaica Willd.) at their ecotone. We hypothesised that soils under Scots pine store more SOC and that tree species composition controls the amount and biochemical composition of organic matter inputs, but does not influence physico-chemical stabilization of SOC. At three locations in Central Spain, we assessed SOC stocks in the forest floor and down to 50 cm in the mineral in pure and mixed stands of Pyrenean oak and Scots pine, as well as litterfall inputs over approximately 3 years at two sites. The relative SOC stability in the topsoil (0-10 cm) was determined through size-fractionation (53 μm) into mineral-associated and particulate organic matter and through KMnO₄-reactive C and soil C:N ratio.Scots pine soils stored 95-140 Mg ha⁻¹ of C (forest floor plus 50 cm mineral soil), roughly the double than Pyrenean oak soils (40-80 Mg ha⁻¹ of C), with stocks closely correlated to litterfall rates. Differences were most pronounced in the forest floor and uppermost 10 cm of the mineral soil, but remained evident in the deeper layers. Biochemical indicators of soil organic matter suggested that biochemical recalcitrance of soil organic matter was higher under pine than under oak, contributing as well to a greater SOC storage under pine. Differences in SOC stocks between tree species were mainly due to the particulate organic matter (not associated to mineral particles). Forest conversion from Pyrenean oak to Scots pine may contribute to enhance soil C sequestration, but only in form of mineral-unprotected soil organic matter.     

Microbial properties and litter and soil nutrients after two prescribed fires in developing savannas in an upland Missouri Ozark Forest

On some landscapes periodic fire may be necessary to develop and maintain oak-dominated savannas. We studied the effects of two annual prescribed burns to determine their effect on microbial activity and soil and litter nutrients 1 year after the last burn. Surface litter and soil from the upper 0–5 cm soil layer in three developing savannas (oak-hickory, Quercus-Carya), oak-hickory-pine (Quercus-Carya-Pinus), and pine (Pinus) were collected one year after the second of two annual prescribed burns. Surface litter was analyzed for nutrients and soil was analyzed for phospholipid fatty acids (PLFAs) and nutrients. Surface litter chemistry differed across the three savannas for potassium (K) and boron (B), being significantly (P < 0.05) higher for unburned forest than for burned forest. Among savannas, only sulfur (S) was higher for the pine savanna and B for the oak-hickory savanna, both were higher for unburned forest than for burned forest. For soil, calcium (Ca) and B differed across savannas, being higher for burned forest than for unburned forest. Among savannas, soil pH, Ca, and B concentrations were higher in soil from burned forest than from unburned forest. Total PLFA differed among savannas, but was not affected by burning treatments. However, the amounts of biomarkers for Gram-positive and Gram-negative bacteria were higher while the amount of biomarker for fungal PLFA was lower for burned forest than for unburned forest. Our results indicate that the two annual prescribed burns moderately affected PLFA microbial community structure and litter and soil nutrient concentrations. However, the long-term effects of fire on these study sites are not known and merit further study.     

Species substitution for carbon storage: Sessile oak versus Corsican pine in France as a case study

Species choice is potentially an important management decision for increasing carbon stocks in forest ecosystems. The substitution of a slow-growing hardwood species (Quercus petraea) by a fast-growing conifer plantation (Pinus nigra subsp. laricio) was studied in central France. Simulations of carbon stocks in tree biomass were conducted using stand growth models Fagacées for sessile oak and PNL for Corsican pine. The changes in soil carbon were assessed using the Century model and data from two European soil monitoring networks: 16 km × 16 km grid and RENECOFOR. Carbon in wood products was assessed with life cycle analysis and lifespan of final products. However, only carbon stocks and their variation were accounted for: effects of energy-consuming materials or fossil fuel substitution are excluded from the analysis. To compare the growth of these two types of forest stands, an important part of the study was to assess the productivity of both species at the same site, using National Forest Inventory data.     

Monitoring the effects of gypsy moth defoliation on forest stand dynamics on Cape Cod,Massachusetts: Sampling intervals and appropriate interpretations

Gypsy moth (Lymantria dispar L.) outbreaks affect stand dynamics and successional patterns in pine–oak woodlands of North America's Atlantic coastal plain. Species-poor overstories are dominated by pitch pine (Pinus rigida), black oak (Quercus velutina) and white oak (Quercus alba). Both oaks are preferred food for gypsy moth, and white oak is the dominant late-successional species. We documented, over a 21-year period, the effects of gypsy moth defoliation on forest stand dynamics at Cape Cod National Seashore, Massachusetts. Our goal was to better understand the influence of gypsy moth defoliation on temporal changes in stand composition. Tree species abundance and frequency were measured in 16 stands with varying abundances of pitch pine and oak. Regression of 2-dimesional rates of change calculated from principal components analysis showed that gypsy moth significantly influenced stand dynamics and successional patterns between 1981 and 1992/1993. Defoliation was virtually absent in the 1990s, and analysis of 1992/1993–2002 rates of change showed topographic slope to be the most important factor influencing stand dynamics during this time. For the period 1981–2002 rates of change analysis failed to identify either factor as influencing changes in stand dynamics. We conclude that the 1980s defoliation events had a transitory effect on upland forest composition, but that there exists the potential for significant long-term impact due to recurrent defoliation episodes. Stand composition should be monitored at intervals which roughly correspond to the time it takes for stands to recover to disturbance events, in the case of those we sampled, approximately 10 years.     

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.     

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.     

Role of topography, canopy structure, and floristics in nest-site selection and nesting success of canopy songbirds

Although oaks (Quercus spp.) have historically dominated many forests in eastern North America, forest composition is changing due to anthropogenic impacts on disturbance regimes. Silvicultural practices, such as partial harvesting, are one component of management to promote oak regeneration. From 2007 to 2009 our research examined nest-site selection and nesting success for a guild of five canopy songbirds in upland mixed-oak forests in southeastern Ohio, USA. We monitored >700 nests across three state forests in both open canopy shelterwood stands harvested to approximately 50% stocking, and closed-canopy mature second-growth. Habitat attributes, including topography, canopy structure, and floristics, were measured at nest sites and random plots ?100 m from nests representing microhabitat available within the territory. Canopy songbirds selected specific topographic microclimates: Eastern Wood-pewees (Contopus virens) nested on xeric ridgetops, Blue-gray Gnatcachers (Polioptila caerulea) favored valleys, and Cerulean Warblers (Dendroica cerulea) preferred productive northeast-facing slopes. Nest sites differed among species in terms of concealment, nest support, topographic position on the slope, and basal area of trees >38 cm dbh. Four of the five focal species selected Quercus alba as the nest substrate more than twice as much as available, and three species avoided Quercus rubra. Daily survival rates of nests were negatively associated with basal area of red oak species (both Quercus velutina and Q. rubra) for several species individually and across the canopy-nesting guild. Additional factors related to success included time of season for Eastern Wood-pewees, nest age for Cerulean Warblers, and concealment and size of the support branch for Scarlet Tanagers (Piranga olivacea). In the long-term management for oak regeneration could benefit canopy songbirds, but our results indicate that white oaks, especially Q. alba, may be preferable to red oak species.     

Responses of oaks and tanoaks to the sudden oak death pathogen after 8 y of monitoring in two coastal California forests

Sudden oak death, caused by Phytophthora ramorum, is widely established in mesic forests of coastal central and northern California. In 2000, we placed 18 plots in two Marin County sites to monitor disease progression in coast live oaks (Quercus agrifolia), California black oaks (Q. kelloggii), and tanoaks (Lithocarpus densiflorus), the species that are most consistently killed by the pathogen in these areas. Through early 2008, the numbers of newly infected trees increased for all species. The infection rate for trees that were asymptomatic in 2000 was 5.0% y⁻¹ for coast live oaks, 4.1% y⁻¹ for black oaks and 10.0% y⁻¹ for tanoaks. Mortality rates were 3.1% y⁻¹ for coast live oaks, 2.4% y⁻¹ for black oaks, and 5.4% y⁻¹ for tanoaks. Mortality not attributed to P. ramorum was 0.54% y⁻¹ for coast live oaks, and 0.75% y⁻¹ for tanoaks. Weibull survival models of trees that were asymptomatic in 2000 provided overall median survival times of 13.7 y for coast live oaks, 13.8 y for black oaks, and 8.8 y for tanoaks. Survival of infected (bleeding) trees declined to 9.7 y for coast live oaks, 6.2 y for black oaks, and 5.8 y for tanoaks. Ambrosia beetle attacks on bleeding trees further reduced modeled survival times by 65–80%, reaffirming the earlier finding that beetle attacks on bleeding cankers considerably reduce survival. Across all plots, the modeled time for 90% of trees that were asymptomatic in 2000 to become infected is 36.5 y for coast live oaks and 15.4 y for tanoaks. There was a trend toward higher infection rates as tree diameter increased. Greater than 90% of living coast live oaks that failed during the study had extensive beetle tunneling at the site of the break. Disease intensity in coast live oaks at the plot level was positively associated with bay laurel (Umbellularia californica) basal area and negatively associated with Pacific madrone (Arbutus menziesii) basal area. This study demonstrates the use of survival modeling to characterize the effects of epidemic disease on different species and to project the future of forests infected with tree pathogens.     

The effects of sudden oak death on foliar moisture content and crown fire potential in tanoak

The introduction of non-native pathogens can have profound effects on forest ecosystems resulting in loss of species, changes in species composition, and altered fuel structure. The introduction of Phytophthora ramorum, the pathogen recognized as causing Sudden Oak Death (SOD), leads to rapid decline and mortality of tanoak (Lithocarpus densiflorus) in forests of coastal California, USA. We tracked foliar moisture content (FMC) of uninfected tanoaks, SOD-infected tanoaks, SOD-killed (dead) tanoaks, and surface litter for 12 months. We found that FMC values differed significantly among the three categories of infection. FMC of uninfected tanoaks averaged 82.3% for the year whereas FMC of infected tanoaks had a lower average of 77.8% (ANOVA, P = 0.04). Dead trees had a significantly lower FMC, averaging 12.3% (ANOVA, P < 0.01) for the year. During fire season (June–September), dead tanoak FMC reached a low of 5.8%, with no significant difference between dead canopy fuels and surface litter (ANOVA, P = 0.44). Application of low FMC values to a crown ignition model results in extremely high canopy base height values to escape crown ignition. Remote estimation of dead FMC using 10-h timelag fuel moisture shows a strong correlation between remote automated weather station (RAWS) 10-h timelag fuel moisture data and the FMC of dead leaves (R² = 0.78, P < 0.01). Results from this study will help refine the decision support tools for fire managers in SOD-affected areas as well as conditions in other forests where diseases and insect epidemics have altered forest canopy fuels.