Sustaining eastern oak forests: Synergistic effects of fire and topography on vegetation and fuels.

Across much of the eastern United States, oak forests are undergoing mesophication as shade-tolerant competitors become more abundant and suppress oak regeneration. Given the historical role of anthropogenic surface fires in promoting oak dominance, prescribed fire has become important in efforts to reverse mesophication and sustain oaks. In 2000 we established the Ohio Hills Fire and Fire Surrogate (FFS) study to examine whether repeated prescribed fire (Fire), mechanical partial harvest (Mech), and their combined application (Mech + Fire) reduced the dominance of subcanopy mesophytic competitors, increased the abundance of large oak-hickory advance regeneration, created a more diverse and productive ground-layer flora, and produced fuel beds more conducive to prescribed fire, reducing the risk of high-severity wildfire. Here we report on the ~20-year effects of treatments on vegetation and fuels and examine the support for interactive effects across a topographic-moisture and energy gradient. In general, we found that Fire and Mech + Fire treatments tended to reverse mesophication while the Mech-only treatment did not. The moderate and occasionally high-intensity fires resulted in effects that were ultimately very similar between the two fire treatments but were modulated by topography with increasing fire severity on drier sites. In particular, we found support for an interaction effect between treatment and topography on forest structure and tree regeneration responses. Fire generally reduced mesophytic tree density in the midstory and sapling strata across all site conditions, while leading to substantial gains in the abundance of large oak-hickory advance regeneration on dry and intermediate landscape positions. Fire also promoted ground-layer diversity and created compositionally distinct communities across all site conditions, primarily through the increased richness of native perennial herbs. However, the fire had limited effects on fine surface fuel loading and increased the loading of large woody fuels, potentially increasing the risk of high-severity wildfire during drought conditions. We conclude that two decades of repeated fires, with and without mechanical density reduction, significantly shifted the trajectory of mesophication across most of the landscape, particularly on dry and intermediate sites, highlighting the capacity of a periodic fire regime to sustain eastern oak forests and promote plant diversity but modulated by topography.

Soil Heating in Fire (SheFire): A model and measurement method for estimating soil heating and effects during wildland fires.

Fire has transformative effects on soil biological, chemical, and physical properties in terrestrial ecosystems around the world. While methods for estimating fire characteristics and associated effects aboveground have progressed in recent decades, there remain major challenges in characterizing soil heating and associated effects belowground. Overcoming these challenges is crucial for understanding how fire influences soil carbon storage, biogeochemical cycling, and ecosystem recovery. In this paper, we present a novel framework for characterizing belowground heating and effects. The framework includes (1) an open-source model to estimate fire-driven soil heating, cooling, and the biotic effects of heating across depths and over time (Soil Heating in Fire model; SheFire) and (2) a simple field method for recording soil temperatures at multiple depths using self-contained temperature sensor and data loggers (i.e., iButtons), installed along a wooden stake inserted into the soil (i.e., an iStake). The iStake overcomes many logistical challenges associated with obtaining temperature profiles using thermocouples. Heating measurements provide inputs to the SheFire model, and modeled soil heating can then be used to derive ecosystem response functions, such as heating effects on microorganisms and tissues. To validate SheFire estimates, we conducted a burn table experiment using iStakes to record temperatures that were in turn used to fit the SheFire model. We then compared SheFire predicted temperatures against measured temperatures at other soil depths. To benchmark iStake measurements against those recorded by thermocouples, we co-located both types of sensors in the burn table experiment. We found that SheFire demonstrated skill in interpolating and extrapolating soil temperatures, with the largest errors occurring at the shallowest depths. We also found that iButton sensors are comparable to thermocouples for recording soil temperatures during fires. Finally, we present a case study using iStakes and SheFire to estimate in situ soil heating during a prescribed fire and demonstrate how observed heating regimes would influence seed and tree root vascular cambium survival at different soil depths. This measurement-modeling framework provides a cutting-edge approach for describing soil temperature regimes (i.e., soil heating) through a soil profile and predicting biological responses.

Prosopis glandulosa persistence is facilitated by differential protection of buds during low- and high-energy fires.

Rangelands worldwide have experienced significant shifts from grass-dominated to woody-plant dominated states over the past century. In North America, these shifts are largely driven by overgrazing and landscape-scale fire suppression. Such shifts reduce productivity for livestock, can have broad-scale impacts to biodiversity, and are often difficult to reverse. Restoring grass dominance often involves restoring fire as an ecological process. However, many resprouting woody plants persist following disturbance, including fire, by resprouting from protected buds, rendering fire ineffective for reducing resprouting woody plant density. Recent research has shown that extreme fire (high-energy fires during periods of water stress) may reduce resprouting capacity. This previous research did not examine whether high-energy fires alone would be sufficient to cause mortality. We created an experimental framework for assessing the "buds-protection-resources" hypothesis of resprouting persistence under different fire energies. In July-August 2018 we exposed 48 individuals of a dominant resprouting woody plant in the region, honey mesquite (Prosopis glandulosa), to two levels of fire energy (high and low) and root crown exposure (exposed vs unexposed) and evaluated resprouting capacity. We censused basal and epicormic resprouts for two years following treatment. Water stress was moderate for several months leading up to fires but low in subsequent years. Epicormic and basal buds were somewhat protected from low- and high-energy fire. However, epicormic buds were protected in very few mesquites subjected to high-energy fires. High-energy fires decreased survival, caused loss of apical dominance, and left residual dead stems, which may increase chances of mortality from future fires. Basal resprout numbers were reduced by high-energy fires, which may have additional implications for long-term mesquite survival. While the buds, protection, and resources components of resprouter persistence all played a role in resprouting, high-energy fire decreased mesquite survival and reduced resprouting. This suggests that high-energy fires affect persistence mechanisms to different extents than low-energy fires. In addition, high-energy fires during normal rainfall can have negative impacts on resprouting capacity; water stress is not a necessary precursor to honey mesquite mortality from high-energy fire.

Differential effects of intra-ventral tegmental area ghrelin and glucagon-like peptide-1 on the stimulatory action of D-amphetamine and cocaine-induced ethanol intake in male Sprague Dawley rats.

In order to further elucidate the role of mesolimbic peptides in the expression of ethanol reward, the present study investigated the effects of ghrelin and glucagon-like peptide-1 (GLP-1) on ethanol intake, in addition to ethanol intake stimulated by systemic d-amphetamine or cocaine treatment. While a number of studies suggest that ghrelin plays an important role in mesolimbic reward, emerging data now indicate that GLP-1 receptor mechanisms inhibit reward signaling, possibly by directly or indirectly inhibiting ghrelinergic activity within the mesolimbic system. In the present study all rats were initially habituated to a 6% ethanol solution. We then demonstrated that intraperitoneal injections of d-amphetamine and cocaine increased ethanol intake compared to the vehicle condition. In subsequent testing we examined the effects of ventral tegmental area (VTA) ghrelin or vehicle paired with a fixed dose of d-amphetamine or vehicle. In separate rats we then investigated the impact of the GLP-1 agonist exendin-4 (Ex-4), injected into the VTA, on ethanol intake alone, or when Ex-4 was co-administered with d-amphetamine or cocaine. Our results indicated that VTA ghrelin significantly increased ethanol intake, and most importantly, potentiated the effect of d-amphetamine and cocaine on ethanol consumption. Conversely, VTA Ex-4 inhibited ethanol intake and antagonized the stimulatory effect of d-amphetamine and cocaine on ethanol consumption. In a final study we further demonstrated that VTA Ex-4 treatment significantly inhibited the combined stimulatory effects of ghrelin paired with d-amphetamine or ghrelin paired with cocaine. Overall our findings are consistent with a critical role for both ghrelin and GLP-1 receptor mechanisms in mesolimbic ethanol reward circuitry. Moreover, our results further suggest that ghrelin and GLP-1 modulate the stimulatory effect of psychostimulants on ethanol intake.

Grass bud responses to fire in a semiarid savanna system.

Increasingly, land managers have attempted to use extreme prescribed fire as a method to address woody plant encroachment in savanna ecosystems. The effect that these fires have on herbaceous vegetation is poorly understood. We experimentally examined immediate (<24 hr) bud response of two dominant graminoids, a C3 caespitose grass, Nassella leucotricha, and a C4 stoloniferous grass, Hilaria belangeri, following fires of varying energy (J/m2) in a semiarid savanna in the Edwards Plateau ecoregion of Texas. Treatments included high- and low-energy fires determined by contrasting fuel loading and a no burn (control) treatment. Belowground axillary buds were counted and their activities classified to determine immediate effects of fire energy on bud activity, dormancy, and mortality. High-energy burns resulted in immediate mortality of N. leucotricha and H. belangeri buds (p < .05). Active buds decreased following high-energy and low-energy burns for both species (p < .05). In contrast, bud activity, dormancy, and mortality remained constant in the control. In the high-energy treatment, 100% (n = 24) of N. leucotricha individuals resprouted while only 25% (n = 24) of H. belangeri individuals resprouted (p < .0001) 3 weeks following treatment application. Bud depths differed between species and may account for this divergence, with average bud depths for N. leucotricha 1.3 cm deeper than H. belangeri (p < .0001). Synthesis and applications: Our results suggest that fire energy directly affects bud activity and mortality through soil heating for these two species. It is imperative to understand how fire energy impacts the bud banks of grasses to better predict grass response to increased use of extreme prescribed fire in land management.

The Wildland Fire Heat Budget-Using Bi-Directional Probes to Measure Sensible Heat Flux and Energy in Surface Fires.

Sensible energy is the primary mode of heat dissipation from combustion in wildland surface fires. However, despite its importance to fire dynamics, smoke transport, and in determining ecological effects, it is not routinely measured. McCaffrey and Heskestad (A robust bidirectional low-velocity probe for flame and fire application. Combustion and Flame 26:125-127, 1976) describe measurements of flame velocity from a bi-directional probe which, when combined with gas temperature measurements, can be used to estimate sensible heat fluxes. In this first field application of bi-directional probes, we describe vertical and horizontal sensible heat fluxes during the RxCADRE experimental surface fires in longleaf pine savanna and open ranges at Eglin Air Force Base, Florida. Flame-front sensible energy is the time-integral of heat flux over a residence time, here defined by the rise in gas temperatures above ambient. Horizontal flow velocities and energies were larger than vertical velocities and energies. Sensible heat flux and energy measurements were coordinated with overhead radiometer measurements from which we estimated fire energy (total energy generated by combustion) under the assumption that 17% of fire energy is radiated. In approximation, horizontal, vertical, and resultant sensible energies averaged 75%, 54%, and 64%, respectively, of fire energy. While promising, measurement challenges remain, including obtaining accurate gas and velocity measurements and capturing three-dimensional flow in the field.

Restoring open canopy pine barrens from the ground up: Repeated burns correspond with increased soil hydraulic conductivity.

Prescribed fire is widely used for ecosystem restoration, yet the mechanisms that determine its effectiveness remain poorly characterized. Because soil hydrology influences ecosystem processes like erosion, runoff, and plant competition, it is important to understand how fire affects soil hydrology. A systematic approach to understanding relationships among vegetation, topography, and fire is needed to advance knowledge of how fire influences soil properties that in turn affect restoration success. Our objective was to characterize relationships among burn severity, vegetation, and soil hydrology in a heterogenous landscape under restoration management. Our study took place in a barrens-forest mosaic with recent prescribed fire history ranging from 0 to 10 burns since 1960, and additional variation in fuel loading, burn severity, vegetation cover, topography, and soils. We measured soil hydraulic conductivity (SHC) during two consecutive years, which represented control, prefire, postfire, and 1-year postfire conditions. Regression tree analysis identified an important threshold effect of antecedent soil moisture on SHC; soils with initial moisture < 13% had lower SHC than soils with initial moisture > 13%. Furthermore, above this threshold, sites with intermediate to high recent burn frequency (4-10 burns) had significantly greater SHC than unburned control sites. High fuel loads associated with brush cutting and piling increased SHC at barrens sites but not brush or pine sites, suggesting an interaction between vegetation cover and fire effects on SHC. At the local hillslope scale, toe-slopes had greater SHC than summits. Our results suggest that repeated prescribed fires of moderate to high frequency may enhance SHC, thereby reducing soil water retention and potentially restoring functional pine barren processes that limit woody plant growth. Prescribed fire may therefore be an important management tool for reversing mesophication and restoring a global array of open canopy ecosystems.

Litter Species Composition and Topographic Effects on Fuels and Modeled Fire Behavior in an Oak-Hickory Forest in the Eastern USA.

Mesophytic species (esp. Acer rubrum) are increasingly replacing oaks (Quercus spp.) in fire-suppressed, deciduous oak-hickory forests of the eastern US. A pivotal hypothesis is that fuel beds derived from mesophytic litter are less likely than beds derived from oak litter to carry a fire and, if they do, are more likely to burn at lower intensities. Species effects, however, are confounded by topographic gradients that affect overstory composition and fuel bed decomposition. To examine the separate and combined effects of litter species composition and topography on surface fuel beds, we conducted a common garden experiment in oak-hickory forests of the Ohio Hills. Each common garden included beds composed of mostly oak and mostly maple litter, representative of oak- and maple-dominated stands, respectively, and a mixture of the two. Beds were replenished each fall for four years. Common gardens (N = 16) were established at four topographic positions (ridges, benches on south- and northeast-facing slopes, and stream terraces) at each of four sites. Litter source and topographic position had largely independent effects on fuel beds and modeled fire dynamics after four years of development. Loading (kg m-2) of the upper litter layer (L), the layer that primarily supports flaming spread, was least in more mesic landscape positions and for maple beds, implying greater decomposition rates for those situations. Bulk density in the L layer (kg m-3) was least for oak beds which, along with higher loading, would promote fire spread and fireline intensity. Loading and bulk density of the combined fermentation and humic (FH) layers were least on stream terrace positions but were not related to species. Litter- and FH-layer moistures during a 5-day dry-down period after a rain event were affected by time and topographic effects while litter source effects were not evident. Characteristics of flaming combustion determined with a cone calorimeter pointed to greater fireline intensity for oak fuel beds and unexpected interactions between litter source and topography. A spread index, which synthesizes a suite of fuel bed, particle, and combustion characteristics to indicate spread (vs extinction) potential, was primarily affected by litter source and, secondarily, by the low spread potentials on mesic landscape positions early in the 5-day dry-down period. A similar result was obtained for modeled fireline intensity. Our results suggest that the continuing transition from oaks to mesophytic species in the Ohio Hills will reduce fire spread potentials and fire intensities.

FireStem2D--a two-dimensional heat transfer model for simulating tree stem injury in fires.

FireStem2D, a software tool for predicting tree stem heating and injury in forest fires, is a physically-based, two-dimensional model of stem thermodynamics that results from heating at the bark surface. It builds on an earlier one-dimensional model (FireStem) and provides improved capabilities for predicting fire-induced mortality and injury before a fire occurs by resolving stem moisture loss, temperatures through the stem, degree of bark charring, and necrotic depth around the stem. We present the results of numerical parameterization and model evaluation experiments for FireStem2D that simulate laboratory stem-heating experiments of 52 tree sections from 25 trees. We also conducted a set of virtual sensitivity analysis experiments to test the effects of unevenness of heating around the stem and with aboveground height using data from two studies: a low-intensity surface fire and a more intense crown fire. The model allows for improved understanding and prediction of the effects of wildland fire on injury and mortality of trees of different species and sizes.