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Plant competition may intensify with climate warming, but whether this will occur equally for conspecific and heterospecific competition remains unknown. Competitive shifts have the potential to instigate community change because the relative strengths of conspecific and heterospecific negative density dependence mediate the stabilizing mechanisms underpinning species coexistence. We examined a mature temperate forest to assess both direct and indirect climate effects at multiple scales: individual species, interspecies relationships, and community stability mechanisms. Our coupled approach (1) quantified tree mortality risk dependence on the interactive effects of competition, climatic water deficit, snowpack, and soil moisture for 28,913 trees over 8 years (3149 mortalities), then (2) used a climate‐projection ensemble to forecast changes in conspecific and heterospecific competition from 2020 to 2100. We predict that projected climate warming will destabilize the foundational forest community by increasing the strength of heterospecific competition at a greater rate and to a greater degree than conspecific competition for four of five abundant tree species, particularly on dry microsites. Modeling showed that these findings were most pronounced after the year 2038, at which point snowpacks were projected to be too small to ameliorate the effects of drought on competitive interactions. Our finding that heterospecific competition is more sensitive than conspecific competition to climate warming may indicate the impending loss of ecosystem functioning. We join the growing body of work showing a predominance of indirect drought effects, yet coupled climate models still fail to consider how changing community dynamics may impact forest cover and, in turn, disrupt forest–climate carbon feedbacks. Ecosystems sharing characteristics with our example forest—those with low species richness and therefore a limited biodiversity insurance effect—may be similarly vulnerable to climate‐mediated destabilization. In such communities, increased heterospecific competition among even a small number of species can more easily destabilize communities without recourse from redundant species. This study of an overlooked but vital mechanism of community change can be adapted by research in a range of ecosystems to improve the understanding of climate change consequences. [ABSTRACT FROM AUTHOR]

Snow duration in post‐fire forests is influenced by neighbourhoods of trees, snags, and deadwood. We used annually resolved, spatially explicit tree and tree mortality data collected in an old‐growth, mixed‐conifer forest in the Sierra Nevada, California, that burned at low to moderate severity to calculate 10 tree neighbourhood metrics for neighbourhoods up to 40 m from snow depth and snow disappearance sampling points. We developed two linear mixed models, predicting snow disappearance timing as a function of tree neighbourhood, litter density, and simulated incoming solar radiation, and two multiple regression models explaining variation in snow depth as a function of tree neighbourhood. Higher densities of post‐fire large‐diameter snags within 10 m of a sampling point were related to higher snow depth (indicating reduced snow interception). Higher densities of large‐diameter trees within 5 m and larger amounts of litter were associated with shorter snow duration (indicating increased longwave radiation emittance and accelerated snow albedo decay). However, live trees with diameters >60 cm within 10 m of a snow disappearance sampling point were associated with a longer‐lasting spring snowpack. This suggests that, despite the local effects of canopy interception and emitted longwave radiation from boles of large trees, shading from their canopies may prolong snow duration over a larger area. Therefore, conservation of widely spaced, large‐diameter trees is important in old‐growth forests because they are resistant to fire and can enhance the seasonal duration of snowmelt. [ABSTRACT FROM AUTHOR]

Background tree mortality is a complex process that requires large sample sizes and long timescales to disentangle the suite of ecological factors that collectively contribute to tree stress, decline, and eventual mortality. Tree mortality associated with acute disturbance events, in contrast, is conspicuous and frequently studied, but there remains a lack of research regarding the role of background mortality processes in mediating the severity and delayed effects of disturbance. We conducted an empirical study by measuring the rates, causes, and spatial pattern of mortality annually among 32,989 individual trees within a large forest demography plot in the Sierra Nevada. We characterized the relationships between background mortality, compound disturbances (fire and drought), and forest spatial structure, and we integrated our findings with a synthesis of the existing literature from around the world to develop a conceptual framework describing the spatio‐temporal signatures of background and disturbance‐related tree mortality. The interactive effects of fire, drought, and background mortality processes altered the rate, spatial structuring, and ecological consequences of mortality. Before fire, spatially non‐random mortality was only evident among small (1 < cm DBH ≤ 10)‐ and medium (10 < cm DBH ≤ 60)‐diameter classes; mortality rates were low (1.7% per yr), and mortality was density‐dependent among small‐diameter trees. Direct fire damage caused the greatest number of moralities (70% of stems ≥1 cm DBH), but the more enduring effects of this disturbance on the demography and spatial pattern of large‐diameter trees occurred during the post‐fire mortality regime. The combined effects of disturbance and biotic mortality agents provoked density‐dependent mortality among large‐diameter (≥60 cm DBH) trees, eliciting a distinct post‐disturbance mortality regime that did not resemble the pattern of either pre‐fire mortality or direct fire effects. The disproportionate ecological significance of the largest trees renders this mortality regime acutely consequential to the long‐term structure and function of forests. [ABSTRACT FROM AUTHOR]

Questions: We examined the spatial patterns of dominant canopy species in a tropical forest to investigate: (a) what is the niche occupancy of canopy species with respect to topographic gradients; (b) what are the dominant ecological processes that explain their distribution; (c) what are the interactions among the most prevalent canopy species; and (d) what are the interactions between canopy species adults and juveniles trees? Location: Rabi permanent CTFS‐ForestGEO plot, Gabon. Methods: We selected the four most abundant canopy species and one timber species. We used Berman's test to determine the effect of three topographic variables on the distribution of each species and univariate analysis to model the spatial pattern of each species using either an inhomogeneous Poisson process or an inhomogeneous Cox process. We also used a bivariate form of the pair correlation function (PCF) to determine the spatial interaction between species and the correlation among conspecific adult and juvenile trees. Results: Four of the five species had aggregated spatial patterns while Lophira alata showed spatial randomness. Most of the variance in the local tree density was explained by within‐population dispersal processes rather than environmental factors. Bivariate PCF tests showed significant segregation between species associations. Two species exhibited aggregation at small distances between young and adult trees, while others showed either complete spatial randomness at small inter‐tree distances or segregation at large distances. Conclusions: This study showed that the spatial pattern in the majority of canopy species was aggregation. Seed dispersal limitation mainly explained the observed aggregation pattern. Habitat filtering, as evidenced by the influence of topographic variables on niche occupancy, marginally, yet significantly, explained this pattern. The different spatial patterns of the principal species permit their coexistence. Spatial segregation among adult and juvenile trees reveals a strong pattern of either species‐specific seed predation or pathogens. [ABSTRACT FROM AUTHOR]

Graphical abstract Highlights • Small (1 m2 ≤ area ≤ 900 m2) unburned patches occupied 5% of the area burned. • Areas with all fire severity classifications contained small unburned patches. • Tree survival was higher when trees were rooted in an unburned patch. • Small tree survival was higher towards the interior of unburned patches. Abstract Fire refugia – the unburned areas within fire perimeters – are important to the survival of many taxa through fire events and the revegetation of post-fire landscapes. Previous work has shown that species use and benefit from small-scale fire refugia (1–1000 m2), but our understanding of where and how fire refugia form is largely limited to the scale of remotely sensed data (i.e., 900 m2 Landsat pixels). To examine the causes and consequences of small fire refugia, we field-mapped all unburned patches ≥1 m2 within a contiguous 25.6 ha forest plot that burned at generally low-to-moderate severity in the 2013 Yosemite Rim Fire, California, USA. Within the Yosemite Forest Dynamics Plot (YFDP), there were 685 unburned patches ≥1 m2, covering a total unburned area of 12,597 m2 (4.9%). Small refugia occurred in all fire severity classifications. Random forest models showed that the proportion of unburned area of 100 m2 grid cells corresponded to pre-fire density and basal area of trees, distance to the nearest stream, and immediate fire mortality, but the relationships were complex and model accuracy was variable. From a pre-fire population of 34,061 total trees ≥1 cm diameter at breast height (1.37 m; DBH) within the plot (1330 trees ha−1), trees of all five of the most common species and those DBH < 30 cm had higher immediate survival rates if their boles were wholly or partially within an unburned patch (P ≤ 0.001). Trees 1 cm ≤ DBH < 10 cm that survived were located closer to the center of the unburned patch than the edge (mean 1.1 m versus 0.6 m; ANOVA; P ≤ 0.001). Four-year survival rates for trees 1 cm ≤ DBH < 10 cm were 58.8% within small refugia and 2.7% in burned areas (P ≤ 0.001). Species richness and the Shannon Diversity Index were associated with unburned quadrats in NMDS ordinations 3 years post-fire. Burn heterogeneity in mixed-conifer forests likely exists at all scales and small refugia contribute to diversity of forest species and structures. Thus, managers may wish to consider scales from 1-m2 to the landscape when designing fuel reduction prescriptions. The partial predictability of refugia location suggests that further work may lead to predictive models of refugial presence that have considerable potential to preserve ecological function or human habitation in fire-frequent forests. [ABSTRACT FROM AUTHOR]

Post-fire tree mortality models are vital tools used by forest land managers to predict fire effects, estimate delayed mortality and develop management prescriptions. We evaluated the performance of mortality models within the First Order Fire Effects Model (FOFEM) software, and compared their performance to locally-parameterised models based on five different forms. We evaluated all models at the individual tree and stand levels with a dataset comprising 34 174 trees from a mixed-conifer forest in the Sierra Nevada, California that burned in the 2013 Rim Fire. We compared stand-level accuracy across a range of spatial scales, and we used point pattern analysis to test the accuracy with which mortality models predict post-fire tree spatial pattern. FOFEM under-predicted mortality for the three conifers, possibly because of the timing of the Rim Fire during a severe multi-year drought. Locally-parameterised models based on crown scorch were most accurate in predicting individual tree mortality, but tree diameter-based models were more accurate at the stand level for Abies concolor and large-diameter Pinus lambertiana , the most abundant trees in this forest. Stand-level accuracy was reduced by spatially correlated error at small spatial scales, but stabilised at scales ≥1 ha. The predictive error of FOFEM generated inaccurate predictions of post-fire spatial pattern at small scales, and this error could be reduced by improving FOFEM model accuracy for small trees. We conducted an evaluation of post-fire tree mortality models with the largest sample to date, and we summarised model accuracy at multiple scales. We found that First Order Fire Effects Models generally had high accuracy for gymnosperms, but accuracy was poor for angiosperms. Fire-related mortality was elevated because of severe multi-year drought, and we provide new mortality models with improved accuracy for fires that occur during drought. [ABSTRACT FROM AUTHOR]

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 largediameter richness was associated with large-diameter trees being individuals of more common species (r2=.17, p5.002). 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. [ABSTRACT FROM AUTHOR]

Abstract: Questions: What drives individual species–area relationships in temperate coniferous forests? Location: Two 25.6‐ha forest plots on the Pacific Slope of North America, one in California, and one in Washington State. Methods: We mapped all trees ≥1 cm in diameter and examined tree species diversity of their local neighbourhoods by calculating the individual species–area relationship for each species and for each of three diameter classes (saplings, mature trees and large‐diameter trees). Results: In the California plot, small trees in four of the five major species occurred in neighbourhoods with higher levels of diversity than would be expected at random. In the Washington plot, small trees for four of five abundant species had neighbourhoods with lower than expected diversity at distances ≤5 m for small trees. However, at distances >5 m, all five species showed higher than expected diversity in their neighbourhoods. Larger trees at both plots tended to occur in neighbourhoods with lower than expected diversity, and no large‐diameter focal species had neighbourhoods with higher than expected diversity. Conclusion: Diversity and co‐existence in temperate conifer‐dominated forests do not appear to be the result of random processes. Competitive interactions appear to dominate for the largest trees of most species, resulting in neighbourhoods with lower diversity. For smaller trees, we suggest that a positive response to environmental heterogeneity is the likely driver of neighbourhoods with higher than expected diversity, although we cannot rule out the possibility that facilitation or conspecific negative density dependence (CNDD) also play a role. [ABSTRACT FROM AUTHOR]

• Disturbance severity and snow duration have species-specific effects on seedlings. • Lower substrate burn severity disadvantaged Abies concolor seedlings. • Earlier snowmelt in the germination year decreased Abies concolor survival. • Post-fire compositional shifts toward drought-tolerant Pinus species are possible. Climate change is altering forest composition through species-specific responses to fire and drought. Future forest composition will depend on how the different regeneration niches of co-occurring species align with current environmental conditions, especially after fire, which can promote germination by exposing mineral soil. Few studies, however, have examined the effects of disturbance severity and microclimate on post-fire regeneration to define and compare the regeneration niches of co-occurring tree species. We used seven years of annual demography and microenvironment data from a 25.6-ha fully censused, stem-mapped forest dynamics plot in California, USA, to examine how disturbance severity, snow duration, and temperature extremes affect the survival of Abies concolor and Pinus lambertiana seedlings that germinated naturally after a low- to moderate-severity fire. We defined disturbance severity at the microsite level, based on characteristics of the substrate, and at the neighborhood level, based on tree mortality. Both disturbance severity and snow duration had species-specific effects on seedling survival, but these differed by life stage. During the germination year, later snow disappearance was associated with a 0.5 increase in survival probability for A. concolor but hardly affected P. lambertiana ; in contrast, higher neighborhood disturbance severity increased survival of both species. After the germination year, higher substrate burn severity was associated with a 0.8 increase in survival probability for A. concolor but hardly affected P. lambertiana ; higher neighborhood disturbance severity and later snow disappearance increased annual survival of both species, but maximum summer temperature had minimal effect. Overall, available seed, higher substrate burn severity, higher neighborhood disturbance severity, and later snow disappearance promoted natural regeneration. However, lower substrate burn severity and earlier snow disappearance in the germination year disadvantaged A. concolor seedlings, increasing the relative abundance of P. lambertiana seedlings compared to the local tree population. Our results indicate that natural post-fire compositional shifts toward drought-tolerant Pinus species–and away from less drought-tolerant Abies species–are possible in the Sierra Nevada, with potential benefits for forest persistence under climate change. Broadly, we show that species differences in regeneration niches shape how disturbance severity and microclimate affect forest species composition. [ABSTRACT FROM AUTHOR]

• Seedlings were highly aggregated 0 – 2.5 m around deadwood and mature adult trees. • Distance to deadwood was the most influential factor for Pinus flexilis seedlings. • Distance to mature Pinus longaeva was highly influential for P. longaeva seedlings. • Mature trees had similar habitat requirements and spatial patterns as seedlings. The spatial patterns of trees are a defining feature of forests, yet because of the long lifespan of trees, the full origins of these spatial patterns remain unclear. We used 15.6 ha of a mapped forest plot to assess the local habitat characteristics and spatial structure of 1,550 naturally established seedlings and 1,955 adults of Pinus flexilis (limber pine) and Pinus longaeva (Great Basin bristlecone pine) in southern Utah, USA. For both species, proximity to deadwood (large end diameter ≥ 10 cm) and mature trees were key parameters of habitat, likely indicating abiotic or biotic facilitation. Both seedlings and mature trees were highly aggregated 0.0–2.5 m around large pieces of deadwood, which can persist in this ecosystem for centuries or millennia. Seedlings were most often located around deadwood 15–22 cm in diameter, though deadwood size had little influence on the distance at which seedlings established. Seedlings of both P. flexilis and P. longaeva were abundant near heterospecific adult Pinus. The elevational distribution of seedlings and adults of both species were not different, indicating a regeneration niche similar to the niche of mature trees and also that changing climate at this site has not yet affected species distributions. Low amounts of manganese and higher levels of pH and potassium in the mineral soil were associated with P. longaeva seedlings but were less important for P. flexilis. Edaphic patterns combined with the presence of deadwood define the overlap between seedling and mature species habitat and identify the key drivers influencing the spatial structure in this high-elevation, old-growth forest. Likely, persistence of these two Pinus species as well as regeneration or site colonization is more favorable in the presence of large deadwood. [ABSTRACT FROM AUTHOR]

• Density reduction from fire or mechanical damage increased post-fire growth rates. • Density reduction from bark beetle mortality did not affect post-fire growth rates. • Pinus had a stronger post-fire growth response to competitive release than Abies. • Mortality and diameter distribution, not growth, control composition of large trees. Global increases in large-tree mortality and multiple disturbances such as fire, drought, and pestilence increase the importance of understanding how large-tree growth responds to changes in competitive neighborhoods. We used 34,175 spatially mapped trees within 25.6 ha that burned at low to moderate severity in 2013 to investigate how changes to tree neighborhoods affected growth of 3,652 surviving trees of two species, Abies concolor (white fir) and Pinus lambertiana (sugar pine). Using diameter measurements taken in 2014 and 2019, we estimated post-fire radial growth of each surviving tree. We modeled annual basal area increment as a function of diameter and extracted residuals to yield relative growth rate (RGR). We used a multi-model approach and AIC to compare the species-specific effects on RGR of all post-fire tree mortality within a neighborhood radius versus mortality broken down by cause. To represent density reduction around each surviving tree, we computed crowding metrics for neighborhood radii up to 20 m for all tree mortality combined and for each mortality cause. The best-fit model included terms for density reduction within 10 m of the surviving tree due to 1) fire or mechanical damage and 2) bark beetle mortality, as well as their interactions with species and diameter. Density reduction due to fire or mechanical damage had a positive effect on RGR that was nearly twice as strong for P. lambertiana as A. concolor. Tree diameter interacted with density reduction for just P. lambertiana. Small-diameter (10-cm) P. lambertiana had a fourfold larger increase in basal area increment than large-diameter (60-cm) P. lambertiana as density reduction due to fire or mechanical damage ranged from 0 to 100 %. The strong growth response of medium-diameter P. lambertiana to density reductions could eventually increase the abundance of large-diameter (≥60 cm) P. lambertiana. In conjunction with mortality rates, which were lower for large-diameter P. lambertiana (0.97 %) than A. concolor (3.31 %), this could increase the relative abundance of large-diameter P. lambertiana from 51.2 % in 2020 to 62.6 % by 2050. Changes to competitive neighborhoods can have species-specific effects on large-tree growth, an important consideration for long-term management of fire-prone forests. [ABSTRACT FROM AUTHOR]

Foundation species define and structure ecological communities but are difficult to identify before they are declining. Yet, their defining role in ecosystems suggests they should be a high priority for protection and management while they are still common and abundant. We used comparative analyses of six large forest dynamics plots spanning a temperate-to-tropical gradient in the Western Hemisphere to identify statistical "fingerprints" of potential foundation species based on their size-frequency and abundance-diameter distributions, and their spatial association with five measures of diversity of associated woody plant species. Potential foundation species are outliers from the common "reverse-J" size-frequency distribution, and have negative effects on alpha diversity and positive effects on beta diversity at most spatial lags and directions. Potential foundation species also are more likely in temperate forests, but foundational species groups may occur in tropical forests. As foundation species (or species groups) decline, associated landscape-scale (beta) diversity is likely to decline along with them. Preservation of this component of biodiversity may be the most important consequence of protecting foundation species while they are still common. [ABSTRACT FROM AUTHOR]

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 (>50,000 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. Image 1 • Various spectral indices detect different aspects of fire-related tree mortality. • dNBR was not the best index, but it was well suited for general use. • Relativized versions of dNBR did not consistently improve performance. • Range in observed mortality was as high as ±40% around the predicted mean. [ABSTRACT FROM AUTHOR]