Big Fires, Big Trees, and Big Plots: Enhancing our Ecological Understanding of Fire with Unprecedented Field Data

Wildfire is an inexorable process in western landscapes, posing a major challenge to land managers: how can we use fire to restore healthy forests without jeopardizing human communities? The purpose of this dissertation is to produce research that will help guide management and support effective wildland fire use in fire-prone forests. I utilized a longitudinal dataset from a single, large forest plot that burned under serendipitous circumstances during the 2013 Rim Fire. My research revealed that post-fire mortality models under-predict mortality of large trees, and may need to be re-calibrated to perform well under future climates. I used satellite-derived data to estimate fire severity, and found that while severity maps may be accurate at broad scales, they failed to capture fine-scale patterns in fire effects. I examined the spatial elements of fire-related mortality, and demonstrate that beetles, pathogens, and inter-tree competition mediated fire effects and provoked complex, spatially structured mortality for years following fire. Finally, I disentangled the interactive effects of fire, beetles, and drought to provide a more mechanistic understanding of compound disturbance dynamics. This represents the first collection of fire ecology research to emerge from a single, exhaustively sampled, longitudinal monitoring plot. This dissertation not only enhances our ecological understanding of fire, it demonstrates the profound potential for large-scale observational research to contribute novel perspectives to the field of fire science

The Utah Forest Dynamics Plot: Long-Term Ecological Monitoring and Theoretical Ecology in a High-Elevation Subalpine Environment

The Unified Neutral Theory of Biodiversity has been advanced as a universal theory for species coexistence in forests worldwide, but few studies have examined its relevance to high-elevation, stressful environments. I established the Utah Forest Dynamics Plot (UFDP) in a heterogeneous subalpine forest at 3,091 m elevation on the Colorado Plateau to examine three underlying assumptions of neutral theory (functional equivalence, ecological equivalence, and habitat generality) and one prediction (the species abundance distribution). The UFDP comprises 27,845 stems ≥1 cm diameter at breast height of 17 species, 10 genera, and 6 families over 13.6 ha. The neutral model was a poor fit to the observed species abundance distribution, but I did not find the alternative lognormal model to provide a better fit. Using spatial pattern analyses of tree data, topography, and soil type, I found some limited support for the neutral theory assumptions of functional and ecological equivalency, with notable exceptions. Populus tremuloides, Pinus flexilis, and Pinus longaeva were characterized by non-neutral recruitment processes, and Abies bifolia and Populus tremuloides exhibited asymmetric competitive and facilitative interactions. The assumption of habitat generality was strongly contradicted, with all ten abundant species in the UFDP having habitat preference. In this subalpine temperate forest, species diversity and community structure are influenced more by habitat heterogeneity, species differences, and niche selection, with neutral processes playing a lesser role

Detecting Tree Mortality with Landsat-Derived Spectral Indices: Improving Ecological Accuracy by Examining Uncertainty

Satellite-derived fire severity metrics are a foundational tool used to estimate fire effects at the landscape scale. Changes in surface characteristics permit reasonably accurate delineation between burned and unburned areas, but variability in severity within burned areas is much more challenging to detect. Previous studies have relied primarily on categorical data to calibrate severity indices in terms of classification accuracy, but this approach does not readily translate into an expected amount of error in terms of actual tree mortality. We addressed this issue by examining a dataset of 40,370 geolocated trees that burned in the 2013 California Rim Fire using 36 Landsat-derived burn severity indices. The differenced Normalized Burn Ratio (dNBR) performed reliably well, but the differenced SWIR:NIR ratio most accurately predicted percent basal area mortality and the differenced normalized vegetation index (dNDVI) most accurately predicted percent mortality of stems ≥10 cm diameter at breast height. Relativized versions of dNBR did not consistently improve accuracy; the relativized burn ratio (RBR) was generally equivalent to dNBR while RdNBR had consistently lower accuracy. There was a high degree of variability in observed tree mortality, especially at intermediate spectral index values. This translated into a considerable amount of uncertainty at the landscape scale, with an expected range in estimated percent basal area mortality greater than 37% for half of the area burned (\u3e50,000 ha). In other words, a 37% range in predicted mortality rate was insufficient to capture the observed mortality rate for half of the area burned. Uncertainty was even greater for percent stem mortality, with half of the area burned exceeding a 46% range in predicted mortality rate. The high degree of uncertainty in tree mortality that we observed challenges the confidence with which Landsat-derived spectral indices have been used to measure fire effects, and this has broad implications for research and management related to post-fire landscape complexity, distribution of seed sources, or persistence of fire refugia. We suggest ways to account for uncertainty that will facilitate a more nuanced and ecologically-accurate interpretation of fire effects. This study makes three key contributions to the field of remote sensing of fire effects: 1) we conducted the most comprehensive comparison to date of all previously published severity indices using the largest contiguous set of georeferenced tree mortality field data and revealed that the accuracy of both absolute and relative spectral indices depends on the tree mortality metric of interest; 2) we conducted this study in a single, large fire that enabled us to isolate variability due to intrinsic, within-landscape factors without the additional variance due to extrinsic factors associated with different biogeographies or climatic conditions; and 3) we identified the range in tree mortality that may be indistinguishable based on spectral indices derived from Landsat satellites, and we demonstrated how this variability translates into a considerable amount of uncertainty in fire effects at the landscape scale

Fuel Dynamics After Reintroduced Fire in an Old-Growth Sierra Nevada Mixed-Conifer Forest

Background: Surface fuel loadings are some of the most important factors contributing to fire intensity and fire spread. In old-growth forests where fire has been long excluded, surface fuel loadings can be high and can include woody debris ≥100 cm in diameter. We assessed surface fuel loadings in a long-unburned old-growth mixed-conifer forest in Yosemite National Park, California, USA, and assessed fuel consumption from a management-ignited fire set to control the progression of the 2013 Rim Fire. Specifically, we characterized the distribution and heterogeneity of pre-fire fuel loadings, both along transects and contained in duff mounds around large trees. We compared surface fuel consumption to that predicted by the standard First Order Fire Effects Model (FOFEM) based on pre-fire fuel loadings and fuel moistures. We also assessed the relationship between tree basal area—calculated for two different spatial neighborhood scales—and pre-fire fuel loadings. Results: Pre-fire total surface fuel loading averaged 192 Mg ha−1 and was reduced by 79% by the fire to 41 Mg ha−1 immediately after fire. Most fuel components were reduced by 87% to 90% by the fire, with the exception of coarse woody debris (CWD), which was reduced by 60%. Litter depth in duff mounds were within 1 SD of plot means, but duff biomass for the largest trees (\u3e150 cm diameter at breast height [DBH]) exceeded plot background levels. Overstory basal area generally had significant positive relationships with pre-fire fuel loadings of litter, duff, 1-hour, and 10-hour fuels, but the strength of the relationships differed between overstory components (live, dead, all [live and dead], species), and negative relationships were observed between live Pinus lambertiana Douglas basal area and CWD. FOFEM over-predicted rotten CWD consumption and under-predicted duff consumption. Conclusions: Surface fuel loadings were characterized by heterogeneity and the presence of large pieces. This heterogeneity likely contributed to differential fire behavior at small scales and heterogeneity in the post-fire environment. The reductions in fuel loadings at our research site were in line with ecological restoration objectives; thus, ecologically restorative burning during fire suppression is possible

Genetic and Spatial Structuring of Populus tremuloides in a Mixed-Species Forest of Southwestern Utah, USA

Populus tremuloides Michx. (aspen) is an iconic species of the southwestern United States, where it is known for its extensive clonality. The size of clones and pattern of clonal distribution within and among stands can provide important clues to the species\u27 evolution and ecology, but there are very few studies that have conducted the type of sampling necessary to define these features. We examined the genetic composition and habitat associations of aspen in a mixed-species forest in Cedar Breaks National Monument on the Markagunt Plateau, southwestern Utah. Genetic analysis of 94 stems ≥1 cm diameter at breast height (dbh) selected from a population census of 2742 stems within a contiguous 13.64-ha plot revealed 2 spatially cohesive triploid genets and 2 diploid genets (all differing in 8 to 15 alleles). Aspen abundance within the 13.64 ha varied between 0 and 634 stems/ha across 8 distinct habitat types. Regenerating aspen stems (1 cm ≤ dbh \u3c 5 cm) varied between 0 and 112 stems/ha, with higher levels of regeneration in habitats with greater aspen dominance relative to other tree species. Recent regeneration may have been stimulated by a Dendroctonous rufipennis outbreak in the 1990s, which killed a high proportion of Picea engelmannii. Even though the visual impression is of a single aspen clone, the 4 identified genets suggest a higher-than-expected level of genetic diversity in this mixed-species stand which may confer resilience to increasing climate variability and drought. Furthermore, aspen regeneration in areas of both low and high adult aspen densities show that these mixed stands can support vigorous aspen populations

Spatially nonrandom tree mortality and ingrowth maintain equilibrium pattern in an old-growth \u3ci\u3ePseudotsuga–Tsuga\u3c/i\u3e forest

Mortality processes in old-growth forests are generally assumed to be driven by gap-scale disturbance, with only a limited role ascribed to density-dependent mortality, but these assumptions are rarely tested with data sets incorporating repeated measurements. Using a 12-ha spatially explicit plot censused 13 years apart in an approximately 500-year-old Pseudotsuga–Tsuga forest, we demonstrate significant density-dependent mortality and spatially aggregated tree recruitment. However, the combined effect of these strongly nonrandom demographic processes was to maintain tree patterns in a state of dynamic equilibrium. Density-dependent mortality was most pronounced for the dominant latesuccessional species, Tsuga heterophylla. The long-lived, early-seral Pseudotsuga menziesii experienced an annual stem mortality rate of 0.84% and no new recruitment. Late-seral species Tsuga and Abies amabilis had nearly balanced demographic rates of ingrowth and mortality. The 2.34% mortality rate for Taxus brevifolia was higher than expected, notably less than ingrowth, and strongly affected by proximity to Tsuga. Large-diameter Tsuga structured both the regenerating conspecific and heterospecific cohorts with recruitment of Tsuga and Abies unlikely in neighborhoods crowded with large-diameter competitors (P , 0.001). Densitydependent competitive interactions strongly shape forest communities even five centuries after stand initiation, underscoring the dynamic nature of even equilibrial old-growth forests

Shrub Communities, Spatial Patterns, and Shrub-Mediated Tree Mortality following Reintroduced Fire in Yosemite National Park, California, USA

Shrubs contribute to the forest fuel load; their distribution is important to tree mortality and regeneration, and vertebrate occupancy. We used a method new to fire ecology—extensive continuous mapping of trees and shrub patches within a single large (25.6 ha) study site—to identify changes in shrub area, biomass, and spatial pattern due to fire reintroduction by a backfire following a century of fire exclusion in lower montane forests of the Sierra Nevada, California, USA. We examined whether trees in close proximity to shrubs prior to fire experienced higher mortality rates than trees in areas without shrubs. We calculated shrub biomass using demography subplots and existing allometric equations, and we developed new equations for beaked hazel (Corylus cornuta ssp. californica [A. de Candolle] E. Murray) from full dissection of 50 stems. Fire decreased shrub patch area from 15.1 % to 0.9 %, reduced live shrub biomass from 3.49 Mg ha−1 to 0.27 Mg ha−1, and consumed 4.41 Mg ha−1 of living and dead shrubs. Distinct (non-overlapping) shrub patches decreased from 47 ha−1 to 6 ha−1. The mean distance between shrub patches increased 135 %. Distances between montane chaparral patches increased 285 %, compared to a 54 % increase in distances between riparian shrub patches and an increase of 267 % between generalist shrub patches. Fire-related tree mortality within shrub patches was marginally lower (67.6 % versus 71.8 %), showing a contrasting effect of shrubs on tree mortality between this forest ecosystem and chaparral-dominated ecosystems in which most trees are killed by fire

Large-Diameter Trees Dominate Snag and Surface Biomass Following Reintroduced Fire

The reintroduction of fire to landscapes where it was once common is considered a priority to restore historical forest dynamics, including reducing tree density and decreasing levels of woody biomass on the forest floor. However, reintroducing fire causes tree mortality that can have unintended ecological outcomes related to woody biomass, with potential impacts to fuel accumulation, carbon sequestration, subsequent fire severity, and forest management. In this study, we examine the interplay between fire and carbon dynamics by asking how reintroduced fire impacts fuel accumulation, carbon sequestration, and subsequent fire severity potential. Beginning pre-fire, and continuing 6 years post-fire, we tracked all live, dead, and fallen trees ≥ 1 cm in diameter and mapped all pieces of deadwood (downed woody debris) originating from tree boles ≥ 10 cm diameter and ≥ 1 m in length in 25.6 ha of an Abies concolor/Pinus lambertiana forest in the central Sierra Nevada, California, USA. We also tracked surface fuels along 2240 m of planar transects pre-fire, immediately post-fire, and 6 years post-fire. Six years after moderate-severity fire, deadwood ≥ 10 cm diameter was 73 Mg ha−1, comprised of 32 Mg ha−1 that persisted through fire and 41 Mg ha−1 of newly fallen wood (compared to 72 Mg ha−1 pre-fire). Woody surface fuel loading was spatially heterogeneous, with mass varying almost four orders of magnitude at the scale of 20 m × 20 m quadrats (minimum, 0.1 Mg ha−1; mean, 73 Mg ha−1; maximum, 497 Mg ha−1). Wood from large-diameter trees (≥ 60 cm diameter) comprised 57% of surface fuel in 2019, but was 75% of snag biomass, indicating high contributions to current and future fuel loading. Reintroduction of fire does not consume all large-diameter fuel and generates high levels of surface fuels ≥ 10 cm diameter within 6 years. Repeated fires are needed to reduce surface fuel loading

Response to Comment on “Plant diversity increases with the strength of negative density dependence at the global scale”

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings—(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance—persist even after controlling for other processes that might influence spatial relationships between adults and recruits

Global importance of large-diameter trees

Aim: To examine the contribution of large‐diameter trees to biomass, stand structure, and species richness across forest biomes. Location: Global. Time period: Early 21st century. Major taxa studied: Woody plants. Methods: We examined the contribution of large trees to forest density, richness and biomass using a global network of 48 large (from 2 to 60 ha) forest plots representing 5,601,473 stems across 9,298 species and 210 plant families. This contribution was assessed using three metrics: the largest 1% of trees ≥ 1 cm diameter at breast height (DBH), all trees ≥ 60 cm DBH, and those rank‐ordered largest trees that cumulatively comprise 50% of forest biomass. Results: Averaged across these 48 forest plots, the largest 1% of trees ≥ 1 cm DBH comprised 50% of aboveground live biomass, with hectare‐scale standard deviation of 26%. Trees ≥ 60 cm DBH comprised 41% of aboveground live tree biomass. The size of the largest trees correlated with total forest biomass (r2 = .62, p < .001). Large‐diameter trees in high biomass forests represented far fewer species relative to overall forest richness (r2 = .45, p < .001). Forests with more diverse large‐diameter tree communities were comprised of smaller trees (r2 = .33, p < .001). Lower large‐diameter richness was associated with large‐diameter trees being individuals of more common species (r2 = .17, p = .002). The concentration of biomass in the largest 1% of trees declined with increasing absolute latitude (r2 = .46, p < .001), as did forest density (r2 = .31, p < .001). Forest structural complexity increased with increasing absolute latitude (r2 = .26, p < .001). Main conclusions: Because large‐diameter trees constitute roughly half of the mature forest biomass worldwide, their dynamics and sensitivities to environmental change represent potentially large controls on global forest carbon cycling. We recommend managing forests for conservation of existing large‐diameter trees or those that can soon reach large diameters as a simple way to conserve and potentially enhance ecosystem services