Herbaceous vegetation responses to experimental fire in savannas and forests depend on biome and climate.

Fire-vegetation feedbacks potentially maintain global savanna and forest distributions. Accordingly, vegetation in savanna and forest ecosystems should have differential responses to fire, but fire response data for herbaceous vegetation have yet to be synthesized across biomes. Here, we examined herbaceous vegetation responses to experimental fire at 30 sites spanning four continents. Across a variety of metrics, herbaceous vegetation increased in abundance where fire was applied, with larger responses to fire in wetter and in cooler and/or less seasonal systems. Compared to forests, savannas were associated with a 4.8 (±0.4) times larger difference in herbaceous vegetation abundance for burned versus unburned plots. In particular, grass cover decreased with fire exclusion in savannas, largely via decreases in C4 grass cover, whereas changes in fire frequency had a relatively weak effect on grass cover in forests. These differential responses underscore the importance of fire for maintaining the vegetation structure of savannas and forests.

Predictive accuracy of post-fire conifer death declines over time in models based on crown and bole injury.

A key uncertainty of empirical models of post-fire tree mortality is understanding the drivers of elevated post-fire mortality several years following fire, known as delayed mortality. Delayed mortality can represent a substantial fraction of mortality, particularly for large trees that are a conservation focus in western US coniferous forests. Current post-fire tree mortality models have undergone limited evaluation of how injury level and time since fire interact to influence model accuracy and predictor variable importance. Less severe injuries potentially serve as an indicator for vulnerability to additional stressors such as bark beetle attack or moisture stress. We used a collection of 164,293 individual tree records to examine post-fire tree mortality in eight western USA conifers: Abies concolor, Abies grandis, Calocedrus decurrens, Larix occidentalis, Pinus contorta, Pinus lambertiana, Pinus ponderosa, and Pseudotsuga menziesii. We evaluated the importance of fire injury predictors on discriminating between surviving trees versus immediate and delayed post-fire mortality. We fit balanced random forest models for each species using cumulative tree mortality from 1 to 5-years post-fire. We compared these results to multi-class random forest models using first-year mortality, 2-5-year mortality, and survival 5-years post-fire as a response variable. Crown volume scorched, diameter at breast height, and relative bark char height, were used as predictor variables. The cumulative mortality models all predicted trees that died within 1-year of fire with high accuracy but failed to predict 2-5-year mortality. The multi-class models were an improvement but had lower accuracy for predicting 2-5-year mortality. Multi-class model accuracies ranged from 85% to 95% across all species for predicting 1-year post-fire mortality, 42%-71% for predicting 2-5-year mortality, and 64%-85% for predicting trees that lived past 5-years. Our study highlights the differences in tree species tolerance to fire injury and suggests that including second-order predictors such as beetle attack or climatic water stress before and after fire will be critical to improve accuracy and better understand the mechanisms and patterns of fire-caused tree death. Random forest models have potential for management applications such as post-fire harvesting and simulating future stand dynamics.

Understanding flammability and bark thickness in the genus Pinus using a phylogenetic approach.

Pinus species dominate fire-prone ecosystems throughout the northern hemisphere. Their litter drive fires that control plant community flammability and multiple ecological processes. To better understand the patterns and mechanisms of pine flammability, we measured leaf characteristics (needle length and thickness) and conducted combustion experiments on litter from 31 species. We paired flammability results with bark accumulation data and used phylogenetic generalized least squares regression to examine relationships between physical traits and flammability. Pine flammability varied widely among pines: flame heights and fuel consumption varied three-fold, and flaming and smoldering durations varied three- to six-fold. Subgenus Pinus species were the most flammable and subgenus Strobus species had the lowest flammability. Needle length was the best predictor of flammability with a significant interaction with subgenus, suggesting that flammability of pines in subgenus Strobus was more affected by physical traits than pines in subgenus Pinus. Species in the subgenus Pinus that accumulated outer bark rapidly also had high flammability, while the relationship was not significant in subgenus Strobus. These results highlight the diverse patterns of flammability in North American pines and the complexity in the mechanisms causing differential flammability.

COVID-19 lockdowns drive decline in active fires in southeastern United States.

Fire is a common ecosystem process in forests and grasslands worldwide. Increasingly, ignitions are controlled by human activities either through suppression of wildfires or intentional ignition of prescribed fires. The southeastern United States leads the nation in prescribed fire, burning ca. 80% of the country's extent annually. The COVID-19 pandemic radically changed human behavior as workplaces implemented social-distancing guidelines and provided an opportunity to evaluate relationships between humans and fire as fire management plans were postponed or cancelled. Using active fire data from satellite-based observations, we found that in the southeastern United States, COVID-19 led to a 21% reduction in fire activity compared to the 2003 to 2019 average. The reduction was more pronounced for federally managed lands, up to 41% below average compared to the past 20 y (38% below average compared to the past decade). Declines in fire activity were partly affected by an unusually wet February before the COVID-19 shutdown began in mid-March 2020. Despite the wet spring, the predicted number of active fire detections was still lower than expected, confirming a COVID-19 signal on ignitions. In addition, prescribed fire management statistics reported by US federal agencies confirmed the satellite observations and showed that, following the wet February and before the mid-March COVID-19 shutdown, cumulative burned area was approaching record highs across the region. With fire return intervals in the southeastern United States as frequent as 1 to 2 y, COVID-19 fire impacts will contribute to an increasing backlog in necessary fire management activities, affecting biodiversity and future fire danger.

Tree crown injury from wildland fires: causes, measurement and ecological and physiological consequences.

The dead foliage of scorched crowns is one of the most conspicuous signatures of wildland fires. Globally, crown scorch from fires in savannas, woodlands and forests causes tree stress and death across diverse taxa. The term crown scorch, however, is inconsistently and ambiguously defined in the literature, causing confusion and conflicting interpretation of results. Furthermore, the underlying mechanisms causing foliage death from fire are poorly understood. The consequences of crown scorch - alterations in physiological, biogeochemical and ecological processes and ecosystem recovery pathways - remain largely unexamined. Most research on the topic assumes the mechanism of leaf and bud death is exposure to lethal air temperatures, with few direct measurements of lethal heating thresholds. Notable information gaps include how energy transfer injures and kills leaves and buds, how nutrients, carbohydrates, and hormones respond, and what physiological consequences lead to mortality. We clarify definitions to encourage use of unified terminology for foliage and bud necrosis resulting from fire. We review the current understanding of the physical mechanisms driving foliar injury, discuss the physiological responses, and explore novel ecological consequences of crown injury from fire. From these elements, we propose research needs for the increasingly interdisciplinary study of fire effects.

Decadal changes in fire frequencies shift tree communities and functional traits.

Global change has resulted in chronic shifts in fire regimes. Variability in the sensitivity of tree communities to multi-decadal changes in fire regimes is critical to anticipating shifts in ecosystem structure and function, yet remains poorly understood. Here, we address the overall effects of fire on tree communities and the factors controlling their sensitivity in 29 sites that experienced multi-decadal alterations in fire frequencies in savanna and forest ecosystems across tropical and temperate regions. Fire had a strong overall effect on tree communities, with an average fire frequency (one fire every three years) reducing stem density by 48% and basal area by 53% after 50 years, relative to unburned plots. The largest changes occurred in savanna ecosystems and in sites with strong wet seasons or strong dry seasons, pointing to fire characteristics and species composition as important. Analyses of functional traits highlighted the impact of fire-driven changes in soil nutrients because frequent burning favoured trees with low biomass nitrogen and phosphorus content, and with more efficient nitrogen acquisition through ectomycorrhizal symbioses. Taken together, the response of trees to altered fire frequencies depends both on climatic and vegetation determinants of fire behaviour and tree growth, and the coupling between fire-driven nutrient losses and plant traits.

The Fire and Tree Mortality Database, for empirical modeling of individual tree mortality after fire.

Wildland fires have a multitude of ecological effects in forests, woodlands, and savannas across the globe. A major focus of past research has been on tree mortality from fire, as trees provide a vast range of biological services. We assembled a database of individual-tree records from prescribed fires and wildfires in the United States. The Fire and Tree Mortality (FTM) database includes records from 164,293 individual trees with records of fire injury (crown scorch, bole char, etc.), tree diameter, and either mortality or top-kill up to ten years post-fire. Data span 142 species and 62 genera, from 409 fires occurring from 1981-2016. Additional variables such as insect attack are included when available. The FTM database can be used to evaluate individual fire-caused mortality models for pre-fire planning and post-fire decision support, to develop improved models, and to explore general patterns of individual fire-induced tree death. The database can also be used to identify knowledge gaps that could be addressed in future research.

Comment on "The global tree restoration potential".

Bastin et al's estimate (Reports, 5 July 2019, p. 76) that tree planting for climate change mitigation could sequester 205 gigatonnes of carbon is approximately five times too large. Their analysis inflated soil organic carbon gains, failed to safeguard against warming from trees at high latitudes and elevations, and considered afforestation of savannas, grasslands, and shrublands to be restoration.

Allometry of the pyrophytic Aristida in fire-maintained longleaf pine-wiregrass ecosystems.

PREMISE OF THE STUDY: Aboveground biomass (AGB) of herbaceous vegetation is a primary source of fuel in frequent surface fires that maintain grasslands, savannas, and woodlands. Methods for nondestructively estimating AGB are required to understand the mechanisms by which fuels affect fire behavior and the effects of time since the last burn. We developed allometric equations to estimate AGB in wiregrass (Aristida beyrichiana/A. stricta), a dominant bunchgrass in Pinus palustris ecosystems and a key species for ecological restoration. METHODS: We collected wiregrass from North Carolina to Florida, across a range of time-since-last burn and site types. We tested 32 mixed effect models to see which predictors were best at predicting live, dead, and total AGB. We also examined how time since burn (TSB) affected the live-to-dead ratio (LDR) using regression. KEY RESULTS: Wiregrass AGB was found to increase with increasing latitude (relative to tussock volume), possibly due to an increase in precipitation, and was greater on more fertile clay soils and flatwoods than on sandy soils. The LDR decreased as a power function with TSB, resulting in rapid accumulation of dead, highly flammable, biomass in the fire-free period. CONCLUSIONS: Greater biomass will support fires of higher intensity. Our models can be useful in the parameterization of future physics-based models to predict fire behavior. Understanding the environmental variables that influence the allometry of wiregrass should help increase the precision of AGB estimates and the subsequent effects on fire behavior and effects on neighboring vegetation.

Linking ecosystem function and hydrologic regime to inform restoration of a forested peatland.

Drainage is a globally common disturbance in forested peatlands that impacts peat soils, forest communities, and associated ecosystem functions, calling for informed hydrologic restoration strategies. The Great Dismal Swamp (GDS), located in Virginia and North Carolina, U.S.A., has been altered since colonial times, particularly by extensive ditch networks installed to lower water levels and facilitate timber harvests. Consequently, peat decomposition rates have accelerated, and red maple has become the dominant tree species, reducing the historical mosaic of bald cypress, Atlantic white-cedar, and pocosin stands. Recent repair and installation of water control structures aim to control drainage and, in doing so, enhance forest community composition and preserve peat depths. To help inform these actions, we established five transects of 15 plots each (75 plots total) along a hydrologic gradient where we measured continuous water levels and ecosystem attributes, including peat depths, microtopography, and forest composition and structure. We found significant differences among transects, with wetter sites having thicker peat, lower red maple importance, greater tree density, and higher overall stand richness. Plot-level analyses comported with these trends, clearly grouping plots by transects (via nonmetric multidimensional scaling) and resulting in significant correlations between specific hydrologic metrics and ecosystem attributes. Our findings highlight hydrologic controls on soil carbon storage, forest structure, and maple dominance, with implications for large-scale hydrologic restoration at GDS and in other degraded forested peatlands more broadly.

Altered community flammability in Florida's Apalachicola ravines and implications for the persistence of the endangered conifer Torreya taxifolia.

Plant species and communities often reflect historic fire regimes via ecological and evolutionary responses to recurrent fires. Plant communities of the southeastern USA experience a wide array of fire regimes, perhaps nowhere more marked than the juxtaposition of fire-prone uplands and adjacent mesic ravines along Florida's Apalachicola River. The ravines contain many endemic and disjunct species, most notably the endangered endemic conifer Torreya taxifolia. A rapid decline in T. taxifolia over the past 60 years has been associated with widespread replacement by other tree species. To understand the changes accompanying the shift in ravine composition, we compared leaf litter flammability of nine historic and contemporary species. We measured maximum flame height, flame duration, smoldering duration, mass loss, absorptive capacity, and drying rate. Ordination and perMANOVA suggest the nine species segregated into three distinct groups: the fire-impeding T. taxifolia and Taxus floridana; an intermediate group of three deciduous angiosperms; and a mixed cluster of four flammable species. Results suggest T. taxifolia and T. floridana were fire-impeding species in these communities, while contemporary dominants burn similarly to the upslope pyric species. The increasing presence of fire-facilitating species may portend a shifting fire regime that further imperils T. taxifolia and other rare species in the formerly fire-safe ravines.

Long-term effects of fire severity on oak-conifer dynamics in the southern Cascades.

We studied vegetation composition and structure in a mixed conifer-oak ecosystem across a range of fire severity 10 years following wildfire. Sample plots centered on focal California black oaks (Quercus kelloggii) were established to evaluate oak and neighboring tree and shrub recovery across a gradient of fire severity in the southern Cascade Range, USA. Shrub and oak resprouting was strongest around focal oaks where conifer mortality was greatest. Linear modeling revealed negative relationships between California black oak sprout height or basal area and residual overstory tree survival, primarily white fir (Abies concolor). The two dominant competing species, California black oak and white fir, showed opposite responses to fire severity. Sprouting California black oak and associated shrubs dominated in severely burned areas, while surviving, non-sprouting white fir maintained dominance by its height advantage and shading effects in areas that burned with low fire severity. Our results indicate that high-severity fire promotes persistence and restoration of ecosystems containing resprouting species, such as California black oak, that are increasingly rare due to widespread fire exclusion in landscapes that historically experienced more frequent fire. We present a conceptual model based on our results and supported by a synthesis of postfire resprouting dynamics literature. Our results and conceptual model help illuminate long-term postfire vegetation responses and the potential ability of fire to catalyze formation of alternate vegetation community structures that may not be apparent in studies that evaluate postfire effects at shorter time-since-fire intervals or at coarser scales.

Unexpected redwood mortality from synergies between wildfire and an emerging infectious disease.

An under-examined component of global change is the alteration of disturbance regimes due to warming climates, continued species invasions, and accelerated land-use change. These drivers of global change are themselves novel ecosystem disturbances that may interact with historically occurring disturbances in complex ways. Here we use the natural experiment presented by wildfires in redwood forests impacted by an emerging infectious disease to demonstrate unexpected synergies of novel disturbance interactions. The dominant tree, coast redwood (fire resistant without negative disease impacts), experienced unexpected synergistic increases in mortality when fire and disease co-occurred. The increased mortality risk, more than fourfold at the peak of the effect, was not predictable from impacts of either disturbance alone. Changes in fire behavior associated with changes to forest fuels that occurred through disease progression overwhelmed redwood's usual resilience to wildfire. Our results demonstrate the potential for interacting disturbances to initiate novel successional trajectories and compromise ecosystem resilience.

Toward a mechanism for eastern North American forest mesophication: differential litter drying across 17 species.

Long-term fire exclusion has altered ecological function in many forested ecosystems in North America. The invasion of fire-sensitive tree species into formerly pyrogenic upland forests in the southeastern United States has resulted in dramatic shifts in surface fuels that have been hypothesized to cause reductions in plant community flammability. The mechanism for the reduced flammability or "mesophication" has lacked empirical study. Here we evaluate a potential mechanism of reduced flammability by quantifying moisture retention (response time and initial moisture capacity) of foliar litter beds from 17 southeastern tree species spanning a wide range of fire tolerance. A k-means cluster analysis resulted in four species groups: a rapidly drying cluster of eight species; a five-species group that absorbed little water but desorbed slowly; a two-species group that absorbed substantial moisture but desorbed rapidly; and a two-species cluster that absorbed substantial moisture and dried slowly. Fire-sensitive species were segregated into the slow moisture loss clusters while fire-tolerant species tended to cluster in the rapid drying groups. Principal-components analysis indicated that several leaf characteristics correlated with absorption capacity and drying rates. Thin-leaved species with high surface area : volume absorbed the greatest moisture content, while those with large, curling leaves had the fastest drying rates. The dramatic shifts in litter fuels as a result of invasion by fire-sensitive species generate a positive feedback that reduce the windows of ignition, thereby facilitating the survival, persistence, and continued invasion of fire-sensitive species in the uplands of the southeastern United States.