Fuel connectivity, burn severity, and seed bank survivorship drive ecosystem transformation in a semiarid shrubland.

A key challenge in ecology is understanding how multiple drivers interact to precipitate persistent vegetation state changes. These state changes may be both precipitated and maintained by disturbances, but predicting whether the state change will be fleeting or persistent requires an understanding of the mechanisms by which disturbance affects the alternative communities. In the sagebrush shrublands of the western United States, widespread annual grass invasion has increased fuel connectivity, which increases the size and spatial contiguity of fires, leading to postfire monocultures of introduced annual grasses (IAG). The novel grassland state can be persistent and is more likely to promote large fires than the shrubland it replaced. But the mechanisms by which prefire invasion and fire occurrence are linked to higher postfire flammability are not fully understood. A natural experiment to explore these interactions presented itself when we arrived in northern Nevada immediately after a 50,000 ha wildfire was extinguished. We hypothesized that the novel grassland state is maintained via a reinforcing feedback where higher fuel connectivity increases burn severity, which subsequently increases postfire IAG dispersal, seed survivorship, and fuel connectivity. We used a Bayesian joint species distribution model and structural equation model framework to assess the strength of the support for each element in this feedback pathway. We found that prefire fuel connectivity increased burn severity and that higher burn severity had mostly positive effects on the occurrence of IAG and another nonnative species and mostly negative or neutral relationships with all other species. Finally, we found that the abundance of IAG seeds in the seed bank immediately after a fire had a positive effect on the fuel connectivity 3 years after the fire, completing a positive feedback promoting IAG. These results demonstrate that the strength of the positive feedback is controlled by measurable characteristics of ecosystem structure, composition, and disturbance. Further, each node in the loop is affected independently by multiple global change drivers. It is possible that these characteristics can be modeled to predict threshold behavior and inform management actions to mitigate or slow the establishment of the grass-fire cycle, perhaps via targeted restoration applications or prefire fuel treatments.

Warming weakens the night-time barrier to global fire.

Night-time provides a critical window for slowing or extinguishing fires owing to the lower temperature and the lower vapour pressure deficit (VPD). However, fire danger is most often assessed based on daytime conditions1,2, capturing what promotes fire spread rather than what impedes fire. Although it is well appreciated that changing daytime weather conditions are exacerbating fire, potential changes in night-time conditions-and their associated role as fire reducers-are less understood. Here we show that night-time fire intensity has increased, which is linked to hotter and drier nights. Our findings are based on global satellite observations of daytime and night-time fire detections and corresponding hourly climate data, from which we determine landcover-specific thresholds of VPD (VPDt), below which fire detections are very rare (less than 95 per cent modelled chance). Globally, daily minimum VPD increased by 25 per cent from 1979 to 2020. Across burnable lands, the annual number of flammable night-time hours-when VPD exceeds VPDt-increased by 110 hours, allowing five additional nights when flammability never ceases. Across nearly one-fifth of burnable lands, flammable nights increased by at least one week across this period. Globally, night fires have become 7.2 per cent more intense from 2003 to 2020, measured via a satellite record. These results reinforce the lack of night-time relief that wildfire suppression teams have experienced in recent years. We expect that continued night-time warming owing to anthropogenic climate change will promote more intense, longer-lasting and larger fires.

Risky Development: Increasing Exposure to Natural Hazards in the United States.

Losses from natural hazards are escalating dramatically, with more properties and critical infrastructure affected each year. Although the magnitude, intensity, and/or frequency of certain hazards has increased, development contributes to this unsustainable trend, as disasters emerge when natural disturbances meet vulnerable assets and populations. To diagnose development patterns leading to increased exposure in the conterminous United States (CONUS), we identified earthquake, flood, hurricane, tornado, and wildfire hazard hotspots, and overlaid them with land use information from the Historical Settlement Data Compilation data set. Our results show that 57% of structures (homes, schools, hospitals, office buildings, etc.) are located in hazard hotspots, which represent only a third of CONUS area, and ∼1.5 million buildings lie in hotspots for two or more hazards. These critical levels of exposure are the legacy of decades of sustained growth and point to our inability, lack of knowledge, or unwillingness to limit development in hazardous zones. Development in these areas is still growing more rapidly than the baseline rates for the nation, portending larger future losses even if the effects of climate change are not considered.

Range dynamics mediated by compensatory life stage responses to experimental climate manipulations.

The expectations of polar or upslope distributional shifts of species ranges in response to warming climate conditions have been recently questioned. Diverse responses of different life stages to changing temperature and moisture regimes may alter these predicted range dynamics. Furthermore, the climate driver(s) influencing demographic rates, and the contribution of each demographic rate to population growth rate (λ), may shift across a species range. We investigated these demographic effects by experimentally manipulating climate and measuring responses of λ in nine populations spanning the elevation range of an alpine plant (Ivesia lycopodioides). Populations exhibited stable growth rates (λ ~ 1) under naturally wet conditions and declining rates (λ < 1) under naturally dry conditions. However, opposing vital rate responses to experimental heating and watering lead to negligible or negative effects on population stability. These findings indicate that life stage-specific responses to changing climate can disrupt the current relationships between population stability and climate across species ranges.

Cross-scale interaction of host tree size and climatic water deficit governs bark beetle-induced tree mortality.

The recent Californian hot drought (2012-2016) precipitated unprecedented ponderosa pine (Pinus ponderosa) mortality, largely attributable to the western pine beetle (Dendroctonus brevicomis; WPB). Broad-scale climate conditions can directly shape tree mortality patterns, but mortality rates respond non-linearly to climate when local-scale forest characteristics influence the behavior of tree-killing bark beetles (e.g., WPB). To test for these cross-scale interactions, we conduct aerial drone surveys at 32 sites along a gradient of climatic water deficit (CWD) spanning 350 km of latitude and 1000 m of elevation in WPB-impacted Sierra Nevada forests. We map, measure, and classify over 450,000 trees within 9 km2, validating measurements with coincident field plots. We find greater size, proportion, and density of ponderosa pine (the WPB host) increase host mortality rates, as does greater CWD. Critically, we find a CWD/host size interaction such that larger trees amplify host mortality rates in hot/dry sites. Management strategies for climate change adaptation should consider how bark beetle disturbances can depend on cross-scale interactions, which challenge our ability to predict and understand patterns of tree mortality.

Local forest structure variability increases resilience to wildfire in dry western U.S. coniferous forests.

A 'resilient' forest endures disturbance and is likely to persist. Resilience to wildfire may arise from feedback between fire behaviour and forest structure in dry forest systems. Frequent fire creates fine-scale variability in forest structure, which may then interrupt fuel continuity and prevent future fires from killing overstorey trees. Testing the generality and scale of this phenomenon is challenging for vast, long-lived forest ecosystems. We quantify forest structural variability and fire severity across >30 years and >1000 wildfires in California's Sierra Nevada. We find that greater variability in forest structure increases resilience by reducing rates of fire-induced tree mortality and that the scale of this effect is local, manifesting at the smallest spatial extent of forest structure tested (90 × 90 m). Resilience of these forests is likely compromised by structural homogenisation from a century of fire suppression, but could be restored with management that increases forest structural variability.

Community turnover by composition and climatic affinity across scales in an alpine system.

PREMISE: Examining community turnover across climate gradients at multiple scales is vital to understanding biogeographic response to climate change. This approach is especially important for alpine plants in which the relative roles of topographic complexity and nonclimatic or stochastic factors vary across spatial scales. METHODS: We examined the structure of alpine plant communities across elevation gradients in the White Mountains, California. Using community climatic niche means (CCNMs) and measures of community dissimilarity, we explored the relation between community composition and elevation gradients at three scales: the mountain range, individual peaks, and within elevation contours. RESULTS: At the mountain range scale, community turnover and CCNMs showed strongly significant relations with elevation, with an increase in the abundance of cooler and wetter-adapted species at higher elevations. At the scale of single peaks, we found weak and inconsistent relations between CCNMs and elevation, but variation in community composition explained by elevation increased. Within the elevation contours, the range of CCNMs was weakly positively correlated with turnover in species identity, likely driven by microclimate and other site-specific factors. CONCLUSIONS: Our results suggest that there is strong environmental sorting of alpine plant communities at broad scales, but microclimatic and site-specific, nonclimatic factors together shape community turnover at finer scales. In the context of climate change, our results imply that community-climate relations are scale-dependent, and predictions of local alpine plant range shifts are limited by a lack of topoclimatic and habitat information.

Parsing propagule pressure: Number, not size, of introductions drives colonization success in a novel environment.

Predicting whether individuals will colonize a novel habitat is of fundamental ecological interest and is crucial to conservation efforts. A consistently supported predictor of colonization success is the number of individuals introduced, also called propagule pressure. Propagule pressure increases with the number of introductions and the number of individuals per introduction (the size of the introduction), but it is unresolved which process is a stronger driver of colonization success. Furthermore, their relative importance may depend upon the environment, with multiple introductions potentially enhancing colonization of fluctuating environments. To evaluate the relative importance of the number and size of introductions and its dependence upon environmental variability, we paired demographic simulations with a microcosm experiment. Using Tribolium flour beetles as a model system, we introduced a fixed number of individuals into replicated novel habitats of stable or fluctuating quality, varying the number of introductions through time and size of each introduction. We evaluated establishment probability and the size of extant populations through seven generations. We found that establishment probability generally increased with more, smaller introductions, but was not affected by biologically realistic fluctuations in environmental quality. Population size was not significantly affected by environmental variability in the simulations, but populations in the microcosms grew larger in a stable environment, especially with more introduction events. In general, the microcosm experiment yielded higher establishment probability and larger populations than the demographic simulations. We suggest that genetic mechanisms likely underlie these differences and thus deserve more attention in efforts to parse propagule pressure. Our results highlight the importance of preventing further introductions of undesirable species to invaded sites and suggest conservation efforts should focus on increasing the number of introductions or reintroductions of desirable species rather than increasing the size of those introduction events into harsh environments.

Effects of topoclimatic complexity on the composition of woody plant communities.

Topography can create substantial environmental variation at fine spatial scales. Shaped by slope, aspect, hill-position and elevation, topoclimate heterogeneity may increase ecological diversity, and act as a spatial buffer for vegetation responding to climate change. Strong links have been observed between climate heterogeneity and species diversity at broader scales, but the importance of topoclimate for woody vegetation across small spatial extents merits closer examination. We established woody vegetation monitoring plots in mixed evergreen-deciduous woodlands that spanned topoclimate gradients of a topographically heterogeneous landscape in northern California. We investigated the association between the structure of adult and regenerating size classes of woody vegetation and multidimensional topoclimate at a fine scale. We found a significant effect of topoclimate on both single-species distributions and community composition. Effects of topoclimate were evident in the regenerating size class for all dominant species (four Quercus spp., Umbellularia californica and Pseudotsuga menziesii) but only in two dominant species (Quercus agrifolia and Quercus garryana) for the adult size class. Adult abundance was correlated with water balance parameters (e.g. climatic water deficit) and recruit abundance was correlated with an interaction between the topoclimate parameters and conspecific adult abundance (likely reflecting local seed dispersal). However, in all cases, the topoclimate signal was weak. The magnitude of environmental variation across our study site may be small relative to the tolerance of long-lived woody species. Dispersal limitations, management practices and patchy disturbance regimes also may interact with topoclimate, weakening its influence on woody vegetation distributions. Our study supports the biological relevance of multidimensional topoclimate for mixed woodland communities, but highlights that this relationship might be mediated by interacting factors at local scales.

Three types of rescue can avert extinction in a changing environment.

Setting aside high-quality large areas of habitat to protect threatened populations is becoming increasingly difficult as humans fragment and degrade the environment. Biologists and managers therefore must determine the best way to shepherd small populations through the dual challenges of reductions in both the number of individuals and genetic variability. By bringing in additional individuals, threatened populations can be increased in size (demographic rescue) or provided with variation to facilitate adaptation and reduce inbreeding (genetic rescue). The relative strengths of demographic and genetic rescue for reducing extinction and increasing growth of threatened populations are untested, and which type of rescue is effective may vary with population size. Using the flour beetle (Tribolium castaneum) in a microcosm experiment, we disentangled the genetic and demographic components of rescue, and compared them with adaptation from standing genetic variation (evolutionary rescue in the strictest sense) using 244 experimental populations founded at either a smaller (50 individuals) or larger (150 individuals) size. Both types of rescue reduced extinction, and those effects were additive. Over the course of six generations, genetic rescue increased population sizes and intrinsic fitness substantially. Both large and small populations showed evidence of being able to adapt from standing genetic variation. Our results support the practice of genetic rescue in facilitating adaptation and reducing inbreeding depression, and suggest that demographic rescue alone may suffice in larger populations even if only moderately inbred individuals are available for addition.