A retrospective assessment of fuel break effectiveness for containing rangeland wildfires in the sagebrush biome.

Escalated wildfire activity within the western U.S. has widespread societal impacts and long-term consequences for the imperiled sagebrush (Artemisia spp.) biome. Shifts from historical fire regimes and the interplay between frequent disturbance and invasive annual grasses may initiate permanent state transitions as wildfire frequency outpaces sagebrush communities' innate capacity to recover. Therefore, wildfire management is at the core of conservation plans for sagebrush ecosystems, especially critical habitat for species of conservation concern such as the greater sage-grouse (Centrocercus urophasianus; hereafter sage-grouse). Fuel breaks help facilitate wildfire suppression by modifying behavior through fuels modification and allowing safe access points for containment by firefighters. The Bureau of Land Management has proposed to roughly double the existing fuel break network in the western U.S., centered on the Great Basin. To our knowledge, no broad-scale examination of fuel break effectiveness or the environmental conditions under which fuel breaks are expected to be most effective has been conducted. We performed a retrospective assessment of probability of fuel break contributing to wildfire containment on recorded wildfire and fuel break interactions from 1985 to 2018 within the western U.S. We characterized environmental, fuels, and weather conditions within 500 m of wildfire contact, and within 5 km of the approaching wildfire. We used a binomial mixed model within a Bayesian framework to identify relationships between these variables and fuel break success. Fuel breaks were least successful in areas classified as having low resilience to disturbance and low resistance to invasion, in areas composed of primarily woody fuels, and when operating in high temperature and low precipitation conditions. Fuel breaks were most effective in areas where fine fuels dominated and in areas that were readily accessible. Maintenance history and fuel break type also contributed to the probability of containment. Overall results indicate a complex and sometimes paradoxical relationship between landscape characteristics that promote wildfire spread and those that impact fuel break effectiveness. Finally, we developed predictive maps of fuel break effectiveness by fuel break type to further elucidate these complex relationships and to inform urgently needed fuel break placement and maintenance priorities across the sagebrush biome.

Integrating anthropogenic factors into regional-scale species distribution models-A novel application in the imperiled sagebrush biome.

Species distribution models (SDMs) that rely on regional-scale environmental variables will play a key role in forecasting species occurrence in the face of climate change. However, in the Anthropocene, a number of local-scale anthropogenic variables, including wildfire history, land-use change, invasive species, and ecological restoration practices can override regional-scale variables to drive patterns of species distribution. Incorporating these human-induced factors into SDMs remains a major research challenge, in part because spatial variability in these factors occurs at fine scales, rendering prediction over regional extents problematic. Here, we used big sagebrush (Artemisia tridentata Nutt.) as a model species to explore whether including human-induced factors improves the fit of the SDM. We applied a Bayesian hurdle spatial approach using 21,753 data points of field-sampled vegetation obtained from the LANDFIRE program to model sagebrush occurrence and cover by incorporating fire history metrics and restoration treatments from 1980 to 2015 throughout the Great Basin of North America. Models including fire attributes and restoration treatments performed better than those including only climate and topographic variables. Number of fires and fire occurrence had the strongest relative effects on big sagebrush occurrence and cover, respectively. The models predicted that the probability of big sagebrush occurrence decreases by 1.2% (95% CI: -6.9%, 0.6%) when one fire occurs and cover decreases by 44.7% (95% CI: -47.9%, -41.3%) if at least one fire occurred over the 36 year period of record. Restoration practices increased the probability of big sagebrush occurrence but had minimal effect on cover. Our results demonstrate the potential value of including disturbance and land management along with climate in models to predict species distributions. As an increasing number of datasets representing land-use history become available, we anticipate that our modeling framework will have broad relevance across a range of biomes and species.

Climate and disturbance influence self-sustaining stand dynamics of aspen (Populus tremuloides) near its range margin.

Species that are primarily seral may form stable (self-sustaining) communities under certain disturbance regimes or environmental conditions, yet such populations may also be particularly vulnerable to ecological change. Aspen (Populus spp.) are generally considered seral throughout the Northern Hemisphere, including P. tremuloides, the most widely distributed tree species in North America. Recent declines in aspen populations have occurred, especially along drought-sensitive margins of its range and where fire exclusion and herbivory have promoted community transition. However, aspen also forms stable stands, and examination of the mechanisms that influence persistence can offer conservation insights, especially where populations are vulnerable to changing climate or altered disturbance dynamics. We sampled tree age and stand characteristics of isolated aspen forests in the arid Great Basin (USA) to determine if (1) aspen communities are more fire-dependent and seral or fire-independent and stable; (2) ungulate browsing inhibits aspen stability; and (3) temporal patterns of vegetative reproduction (i.e., ramet establishment or "suckering") are correlated with climate. Aspen size and age class densities strongly fit negative exponential distributions, whether grouped geographically or by functional type, suggesting landscape-scale persistence. Continuous age distributions and high proportions of recruitment-sized to overstory trees suggest stability at stand scales, with exceptions including stands with higher browsing pressure. Few stands had evidence of fire, and relationships between dead tree size and variability in live tree size suggest a lack of fire dependency. Several 5-yr averaged climate variables and one sea surface temperature index were correlated with aspen ramet establishment densities over time, with strongest relationships occurring ~5 yr prior to establishment year, often followed by inverse relationships ~1 yr after. Indeed, aspen establishment density for a recent 41-yr period was reliably reconstructed using antecedent climate conditions derived from a single drought index. Temporally synchronized aspen ramet establishment across the study region may be due to climate-driven storage of nonstructural carbohydrate reserves in clonal root systems later used for regeneration. Complex regeneration dynamics of these self-sustaining aspen stands, especially sensitivity to climate variability, suggest they may serve as harbingers of ecological change in the arid Great Basin and in other aspen populations near their range margin.

Intraspecific Niche Models for Ponderosa Pine (Pinus ponderosa) Suggest Potential Variability in Population-Level Response to Climate Change.

Unique responses to climate change can occur across intraspecific levels, resulting in individualistic adaptation or movement patterns among populations within a given species. Thus, the need to model potential responses among genetically distinct populations within a species is increasingly recognized. However, predictive models of future distributions are regularly fit at the species level, often because intraspecific variation is unknown or is identified only within limited sample locations. In this study, we considered the role of intraspecific variation to shape the geographic distribution of ponderosa pine (Pinus ponderosa), an ecologically and economically important tree species in North America. Morphological and genetic variation across the distribution of ponderosa pine suggest the need to model intraspecific populations: the two varieties (var. ponderosa and var. scopulorum) and several haplotype groups within each variety have been shown to occupy unique climatic niches, suggesting populations have distinct evolutionary lineages adapted to different environmental conditions. We utilized a recently available, geographically widespread dataset of intraspecific variation (haplotypes) for ponderosa pine and a recently devised lineage distance modeling approach to derive additional, likely intraspecific occurrence locations. We confirmed the relative uniqueness of each haplotype-climate relationship using a niche-overlap analysis, and developed ecological niche models (ENMs) to project the distribution for two varieties and eight haplotypes under future climate forecasts. Future projections of haplotype niche distributions generally revealed greater potential range loss than predicted for the varieties. This difference may reflect intraspecific responses of distinct evolutionary lineages. However, directional trends are generally consistent across intraspecific levels, and include a loss of distributional area and an upward shift in elevation. Our results demonstrate the utility in modeling intraspecific response to changing climate and they inform management and conservation strategies, by identifying haplotypes and geographic areas that may be most at risk, or most secure, under projected climate change.

Methodological considerations of terrestrial laser scanning for vegetation monitoring in the sagebrush steppe.

Terrestrial laser scanning (TLS) provides fast collection of high-definition structural information, making it a valuable field instrument to many monitoring applications. A weakness of TLS collections, especially in vegetation, is the occurrence of unsampled regions in point clouds where the sensor's line-of-sight is blocked by intervening material. This problem, referred to as occlusion, may be mitigated by scanning target areas from several positions, increasing the chance that any given area will fall within the scanner's line-of-sight from at least one position. Because TLS collections are often employed in remote regions where the scope of sampling is limited by logistical factors such as time and battery power, it is important to design field protocols which maximize efficiency and support increased quantity and quality of the data collected. This study informs researchers and practitioners seeking to optimize TLS sampling methods for vegetation monitoring in dryland ecosystems through three analyses. First, we quantify the 2D extent of occluded regions based on the range from single scan positions. Second, we measure the efficacy of additional scan positions on the reduction of 2D occluded regions (area) using progressive configurations of scan positions in 1 ha plots. Third, we test the reproducibility of 3D sampling yielded by a 5-scan/ha sampling methodology using redundant sets of scans. Analyses were performed using measurements at analysis scales of 5 to 50 cm across the 1-ha plots, and we considered plots in grass and shrub-dominated communities separately. In grass-dominated plots, a center-scan configuration and 5 cm pixel size sampled at least 90% of the area up to 18 m away from the scanner. In shrub-dominated plots, sampling at least 90% of the area was only achieved within a distance of 12 m. We found that 3 and 5 scans/ha are needed to sample at least ~ 70% of the total area (1 ha) in the grass and shrub-dominated plots, respectively, using 5 cm pixels to measure sampling presence-absence. The reproducibility of 3D sampling provided by a 5 position scan layout across 1-ha plots was 50% (shrub) and 70% (grass) using a 5-cm voxel size, whereas at the 50-cm voxel scale, reproducibility of sampling was nearly 100% for all plot types. Future studies applying TLS in similar dryland environments for vegetation monitoring may use our results as a guide to efficiently achieve sampling coverage and reproducibility in datasets.

Exploring Climate Niches of Ponderosa Pine (Pinus ponderosa Douglas ex Lawson) Haplotypes in the Western United States: Implications for Evolutionary History and Conservation.

Ponderosa pine (Pinus ponderosa Douglas ex Lawson) occupies montane environments throughout western North America, where it is both an ecologically and economically important tree species. A recent study using mitochondrial DNA analysis demonstrated substantial genetic variation among ponderosa pine populations in the western U.S., identifying 10 haplotypes with unique evolutionary lineages that generally correspond spatially with distributions of the Pacific (P. p. var. ponderosa) and Rocky Mountain (P. p. var. scopulorum) varieties. To elucidate the role of climate in shaping the phylogeographic history of ponderosa pine, we used nonparametric multiplicative regression to develop predictive climate niche models for two varieties and 10 haplotypes and to hindcast potential distribution of the varieties during the last glacial maximum (LGM), ~22,000 yr BP. Our climate niche models performed well for the varieties, but haplotype models were constrained in some cases by small datasets and unmeasured microclimate influences. The models suggest strong relationships between genetic lineages and climate. Particularly evident was the role of seasonal precipitation balance in most models, with winter- and summer-dominated precipitation regimes strongly associated with P. p. vars. ponderosa and scopulorum, respectively. Indeed, where present-day climate niches overlap between the varieties, introgression of two haplotypes also occurs along a steep clinal divide in western Montana. Reconstructed climate niches for the LGM suggest potentially suitable climate existed for the Pacific variety in the California Floristic province, the Great Basin, and Arizona highlands, while suitable climate for the Rocky Mountain variety may have existed across the southwestern interior highlands. These findings underscore potentially unique phylogeographic origins of modern ponderosa pine evolutionary lineages, including potential adaptations to Pleistocene climates associated with discrete temporary glacial refugia. Our predictive climate niche models may inform strategies for further genetic research (e.g., sampling design) and conservation that promotes haplotype compatibility with projected changes in future climate.

Comparing modern and presettlement forest dynamics of a subboreal wilderness: does spruce budworm enhance fire risk?

Insect disturbance is often thought to increase fire risk through enhanced fuel loadings, particularly in coniferous forest ecosystems. Yet insect disturbances also affect successional pathways and landscape structure that interact with fire disturbances (and vice-versa) over longer time scales. We applied a landscape succession and disturbance model (LANDIS-II) to evaluate the relative strength of interactions between spruce budworm (Choristoneura fumiferana) outbreaks and fire disturbances in the Boundary Waters Canoe Area (BWCA) in northern Minnesota (USA). Disturbance interactions were evaluated for two different scenarios: presettlement forests and fire regimes vs. contemporary forests and fire regimes. Forest composition under the contemporary scenario trended toward mixtures of deciduous species (primarily Betula papyrifera and Populus spp.) and shade-tolerant conifers (Picea mariana, Abies balsamea, Thuja occidentalis), with disturbances dominated by a combination of budworm defoliation and high-severity fires. The presettlement scenario retained comparatively more "big pines" (i.e., Pinus strobus, P. resinosa) and tamarack (L. laricina), and experienced less budworm disturbance and a comparatively less-severe fire regime. Spruce budworm disturbance decreased area burned and fire severity under both scenarios when averaged across the entire 300-year simulations. Contrary to past research, area burned and fire severity during outbreak decades were each similar to that observed in non-outbreak decades. Our analyses suggest budworm disturbances within forests of the BWCA have a comparatively weak effect on long-term forest composition due to a combination of characteristics. These include strict host specificity, fine-scaled patchiness created by defoliation damage, and advance regeneration of its primary host, balsam fir (A. balsamea) that allows its host to persist despite repeated disturbances. Understanding the nature of the three-way interaction between budworm, fire, and composition has important ramifications for both fire mitigation strategies and ecosystem restoration initiatives. We conclude that budworm disturbance can partially mitigate long-term future fire risk by periodically reducing live ladder fuel within the mixed forest types of the BWCA but will do little to reverse the compositional trends caused in part by reduced fire rotations.

Historical fire and multidecadal drought as context for piñon-juniper woodland restoration in western Colorado.

Fire is known to structure tree populations, but the role of broad-scale climate variability is less clear. For example, the influence of climatic "teleconnections" (the relationship between oceanic-atmospheric fluctuations and anomalous weather patterns across broad scales) on forest age structure is relatively unexplored. We sampled semiarid piñon-juniper (Pinus edulis-Juniperus osteosperma) woodlands in western Colorado, U.S.A., to test the hypothesis that woodland age structures are shaped by climate, including links to oceanic-atmospheric fluctuations, and by past fires and livestock grazing. Low-severity surface fire was lacking, as fire scars were absent, and did not influence woodland densities, but stand-replacing fires served as long-rotation (>400-600 years), stand-initiating events. Old-growth stands (>300 years old) were found in 75% of plots, consistent with a long fire rotation. Juniper and piñon age structures suggest contrasting responses during the past several centuries to dry and wet episodes linked to the Atlantic Multidecadal Oscillation (AMO) and Pacific Decadal Oscillation (PDO). Juniper density increased slightly during periods of drought, positive (warm) AMO (after approximately 10-year lag), and negative (cool) PDO. In contrast, piñon populations may still be recovering from a long, drought-filled period (AD 1620-1820), with pulses of recovery favored during cool AMO, warm PDO, and above-average moisture periods. Analysis of 20th-century tree establishment and instrumental climate data corroborate the long-term relationships between age structure and climate. After Euro-American settlement (AD 1881), livestock grazing reduced understory grasses and forbs, reducing competition with tree seedlings and facilitating climate-induced increases in piñons. Thus tree populations in these woodlands are in flux, affected by drought and wet periods linked to oceanic-atmospheric variability, Euro-American livestock grazing, and long-rotation, high-severity fires. Reductions in livestock grazing levels may aid ecological restoration efforts. However, given long-term fluctuations in tree density and composition, and expected further drought, thinning or burning to reduce tree populations may be misdirected.