Multiple social and environmental factors affect wildland fire response of full or less-than-full suppression.

Wildland fire incident commanders make wildfire response decisions within an increasingly complex socio-environmental context. Threats to human safety and property, along with public pressures and agency cultures, often lead commanders to emphasize full suppression. However, commanders may use less-than-full suppression to enhance responder safety, reduce firefighting costs, and encourage beneficial effects of fire. This study asks: what management, socioeconomic, environmental, and fire behavior characteristics are associated with full suppression and the less-than-full suppression methods of point-zone protection, confinement/containment, and maintain/monitor? We analyzed incident report data from 374 wildfires in the United States northern Rocky Mountains between 2008 and 2013. Regression models showed that full suppression was most strongly associated with higher housing density and earlier dates in the calendar year, along with non-federal land jurisdiction, regional and national incident management teams, human-caused ignitions, low fire-growth potential, and greater fire size. Interviews with commanders provided decision-making context for these regression results. Future efforts to encourage less-than-full suppression should address the complex management context, in addition to the biophysical context, of fire response.

Peeking under the canopy: anomalously short fire-return intervals alter subalpine forest understory plant communities.

Climate change is driving changes in disturbance regimes world-wide. In forests adapted to infrequent, high-severity fires, recent anomalously short fire-return intervals (FRIs) have resulted in greatly reduced postfire tree regeneration. However, effects on understory plant communities remain unexplored. Understory plant communities were sampled in 31 plot pairs across Greater Yellowstone (Wyoming, USA). Each pair included one plot burned at high severity twice in < 30 yr and one plot burned in the same most recent fire but not burned previously for > 125 yr. Understory communities following short-interval fires were also compared with those following the previous long-interval fire. Species capable of growing in drier conditions and in lower vegetation zones became more abundant and regional differences in plant communities declined following short-interval fire. Dissimilarity between plot pairs increased in mesic settings and decreased with time since fire and postfire winter snowfall. Reduced postfire tree density following short-interval fire rather than FRI per se affected the occurrence of most plant species. Anomalously short FRIs altered understory plant communities in space and time, with some indications of community thermophilization and regional homogenization. These and other shifts in understory plant communities may continue with ongoing changes in climate and fire across temperate forests.

Less fuel for the next fire? Short-interval fire delays forest recovery and interacting drivers amplify effects.

As 21st-century climate and disturbance dynamics depart from historic baselines, ecosystem resilience is uncertain. Multiple drivers are changing simultaneously, and interactions among drivers could amplify ecosystem vulnerability to change. Subalpine forests in Greater Yellowstone (Northern Rocky Mountains, USA) were historically resilient to infrequent (100-300 year), severe fire. We sampled paired short-interval (<30-year) and long-interval (>125-year) post-fire plots most recently burned between 1988 and 2018 to address two questions: (1) How do short-interval fire, climate, topography, and distance to unburned live forest edge interact to affect post-fire forest regeneration? (2) How do forest biomass and fuels vary following short-interval versus long-interval severe fires? Mean post-fire live tree stem density was an order of magnitude lower following short-interval versus long-interval fires (3240 vs. 28,741 stems ha-1 , respectively). Differences between paired plots were amplified at longer distances to live forest edge. Surprisingly, warmer-drier climate was associated with higher seedling densities even after short-interval fire, likely relating to regional variation in serotiny of lodgepole pine (Pinus contorta var. latifolia). Unlike conifers, density of aspen (Populus tremuloides), a deciduous resprouter, increased with short-interval versus long-interval fires (mean 384 vs. 62 stems ha-1 , respectively). Live biomass and canopy fuels remained low nearly 30 years after short-interval fire, in contrast with rapid recovery after long-interval fire, suggesting that future burn severity may be reduced for several decades following reburns. Short-interval plots also had half as much dead woody biomass compared with long-interval plots (60 vs. 121 Mg ha-1 ), primarily due to the absence of large snags. Our results suggest differences in tree regeneration following short-interval versus long-interval fires will be especially pronounced where serotiny was high historically. Propagule limitation will also interact with short-interval fires to diminish tree regeneration but lessen subsequent burn severity. Amplifying driver interactions are likely to threaten forest resilience under expected trajectories of a future fire.

Regeneration strategies and forest resilience to changing fire regimes: Insights from a Goldilocks model.

Disturbances are ubiquitous in ecological systems, and species have evolved a range of strategies to resist or rebound following disturbance. Understanding how the presence and complementarity of regeneration traits will affect community responses to disturbance is increasingly urgent as disturbance regimes shift beyond their historical ranges of variability. We define "disturbance niche" as a species' fitness across a range of disturbance sizes and frequencies that can reflect the fundamental or realized niche, that is, whether the species occurs alone or with other species. We developed a model of intermediate complexity (i.e., a Goldilocks model) to infer the disturbance niche. We parameterized the model for subalpine forests in Yellowstone National Park (USA) adapted to infrequent stand-replacing fires and included the three major tree-regeneration strategies: (1) obligate seeders that rely on ex situ seeding into burned areas (non-serotinous lodgepole pine, Pinus contorta var. latifola), (2) obligate seeders that depend on in situ seedbanks (serotinous lodgepole pine, Pinus contorta var. latifola), and (3) species that can resprout from surviving roots following fire (quaking aspen, Populus tremuloides). Our results showed which regeneration strategies increase or decrease in prevalence as fire rotation declines. Non-serotinous pines were extirpated when fire rotation was below 50 years in a monoculture and 100 years in a mixed forest; serotinous pines were extirpated when fire rotation was below 20 years; and aspen was extirpated when fire rotation fell below 6 years. The fundamental and realized disturbance niches pinpointed the key mechanisms limiting regeneration for each strategy, namely, increasing fire size for non-serotinous pine (ex situ seeders), decreasing fire frequency for serotinous pine (in situ seeders), and interspecific competition for aspen (resprouters). In a mixed forest, the three regeneration strategies were complementary and each dominated at different combinations of fire size and frequency. Consequently, diversity of regeneration strategies enhanced forest resilience to declining fire rotations. Despite its simplicity, our Goldilocks model produced realistic dynamics and could be readily adapted to other disturbance-prone ecosystems to explore the generality of these results. The disturbance niche is a key concept for anticipating community resilience to changing disturbance regimes.

Post-disturbance reorganization of forest ecosystems in a changing world.

Forest ecosystems are strongly impacted by continuing climate change and increasing disturbance activity, but how forest dynamics will respond remains highly uncertain. Here, we argue that a short time window after disturbance (i.e., a discrete event that disrupts prevailing ecosystem structure and composition and releases resources) is pivotal for future forest development. Trees that establish during this reorganization phase can shape forest structure and composition for centuries, providing operational early indications of forest change. While forest change has been fruitfully studied through a lens of resilience, profound ecological changes can be masked by a resilience versus regime shift dichotomy. We present a framework for characterizing the full spectrum of change after disturbance, analyzing forest reorganization along dimensions of forest structure (number, size, and spatial arrangement of trees) and composition (identity and diversity of tree species). We propose four major pathways through which forest cover can persist but reorganize following disturbance: resilience (no change in structure and composition), restructuring (structure changes but composition does not), reassembly (composition changes but structure does not), and replacement (structure and composition both change). Regime shifts occur when vegetation structure and composition are altered so profoundly that the emerging trajectory leads to nonforest. We identify fundamental processes underpinning forest reorganization which, if disrupted, deflect ecosystems away from resilience. To understand and predict forest reorganization, assessing these processes and the traits modulating them is crucial. A new wave of experiments, measurements, and models emphasizing the reorganization phase will further the capacity to anticipate future forest dynamics.

Widespread regeneration failure in forests of Greater Yellowstone under scenarios of future climate and fire.

Changing climate and disturbance regimes are increasingly challenging the resilience of forest ecosystems around the globe. A powerful indicator for the loss of resilience is regeneration failure, that is, the inability of the prevailing tree species to regenerate after disturbance. Regeneration failure can result from the interplay among disturbance changes (e.g., larger and more frequent fires), altered climate conditions (e.g., increased drought), and functional traits (e.g., method of seed dispersal). This complexity makes projections of regeneration failure challenging. Here we applied a novel simulation approach assimilating data-driven fire projections with vegetation responses from process modeling by means of deep neural networks. We (i) quantified the future probability of regeneration failure; (ii) identified spatial hotspots of regeneration failure; and (iii) assessed how current forest types differ in their ability to regenerate under future climate and fire. We focused on the Greater Yellowstone Ecosystem (2.9 × 106  ha of forest) in the Rocky Mountains of the USA, which has experienced large wildfires in the past and is expected to undergo drastic changes in climate and fire in the future. We simulated four climate scenarios until 2100 at a fine spatial grain (100 m). Both wildfire activity and unstocked forest area increased substantially throughout the 21st century in all simulated scenarios. By 2100, between 28% and 59% of the forested area failed to regenerate, indicating considerable loss of resilience. Areas disproportionally at risk occurred where fires are not constrained by topography and in valleys aligned with predominant winds. High-elevation forest types not adapted to fire (i.e., Picea engelmannii-Abies lasiocarpa as well as non-serotinous Pinus contorta var. latifolia forests) were especially vulnerable to regeneration failure. We conclude that changing climate and fire could exceed the resilience of forests in a substantial portion of Greater Yellowstone, with profound implications for carbon, biodiversity, and recreation.

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 propagule doesn't fall far from the tree, especially after short-interval, high-severity fire.

Subalpine forests that historically burned every 100-300 yr are expected to burn more frequently as climate warms, perhaps before trees reach reproductive maturity or produce a serotinous seedbank. Tree regeneration after short-interval (<30-yr) high-severity fire will increasingly rely on seed dispersal from unburned trees, but how dispersal varies with age and structure of surrounding forest is poorly understood. We studied wind dispersal of three conifers (Picea engelmannii, Abies lasiocarpa, and Pinus contorta var. latifolia, which can be serotinous and nonserotinous) after a stand-replacing fire that burned young (≤30 yr) and older (>100 yr) P. contorta forest in Grand Teton National Park (Wyoming, USA). We asked how propagule pressure varied with time since last fire, how seed delivery into burned forest varied with age and structure of live forest edges, what variables explained seed delivery into burned forest, and how spatial patterns of delivery across the burned area could vary with alternate patterns of surrounding live forest age. Seeds were collected in traps along 100-m transects (n = 18) extending from live forest edges of varying age (18, 30, and >100 yr) into areas of recent (2-yr) high-severity fire, and along transects in live forests to measure propagule pressure. Propagule pressure was low in 18-yr-old stands (~8 seeds/m2 ) and similarly greater in 30- and 100-yr-old stands (~32 seeds/m2 ). Mean dispersal distance was lowest from 18-yr-old edges and greatest from >100-yr-old edges. Seed delivery into burned forest declined with increasing distance and increased with height of trees at live forest edges, and was consistently higher for P. contorta than for other conifers. Empirical dispersal kernels revealed that seed delivery from 18-yr-old edges was very low (≤2.4 seeds/m2 ) and concentrated within 10 m of the live edge, whereas seed delivery from >100-yr-old edges was >4.9 seeds/m2 out to 80 m. When extrapolated throughout the burned landscape, estimated seed delivery was low (<49,400 seeds/ha) in >70% of areas that burned in short-interval fire (<30 yr). As fire frequency increases, immaturity risk will be compounded in short-interval fires because seed dispersal from surrounding young trees is limited.

Simulating forest resilience: A review.

AIM: Simulation models are important tools for quantifying the resilience (i.e., persistence under changed environmental conditions) of forest ecosystems to global change. We synthesized the modelling literature on forest resilience, summarizing common models and applications in resilience research, and scrutinizing the implementation of important resilience mechanisms in these models. Models applied to assess resilience are highly diverse, and our goal was to assess how well they account for important resilience mechanisms identified in experimental and empirical research. LOCATION: Global. TIME PERIOD: 1994 to 2019. MAJOR TAXA STUDIED: Trees. METHODS: We reviewed the forest resilience literature using online databases, selecting 119 simulation modelling studies for further analysis. We identified a set of resilience mechanisms from the general resilience literature and analysed models for their representation of these mechanisms. Analyses were grouped by investigated drivers (resilience to what) and responses (resilience of what), as well as by the type of model being used. RESULTS: Models used to study forest resilience varied widely, from analytical approaches to complex landscape simulators. The most commonly addressed questions were associated with resilience of forest cover to fire. Important resilience mechanisms pertaining to regeneration, soil processes, and disturbance legacies were explicitly simulated in only 34 to 46% of the model applications. MAIN CONCLUSIONS: We found a large gap between processes identified as underpinning forest resilience in the theoretical and empirical literature, and those represented in models used to assess forest resilience. Contemporary forest models developed for other goals may be poorly suited for studying forest resilience during an era of accelerating change. Our results highlight the need for a new wave of model development to enhance understanding of and management for resilient forests.

Pervasive shifts in forest dynamics in a changing world.

Forest dynamics arise from the interplay of environmental drivers and disturbances with the demographic processes of recruitment, growth, and mortality, subsequently driving biomass and species composition. However, forest disturbances and subsequent recovery are shifting with global changes in climate and land use, altering these dynamics. Changes in environmental drivers, land use, and disturbance regimes are forcing forests toward younger, shorter stands. Rising carbon dioxide, acclimation, adaptation, and migration can influence these impacts. Recent developments in Earth system models support increasingly realistic simulations of vegetation dynamics. In parallel, emerging remote sensing datasets promise qualitatively new and more abundant data on the underlying processes and consequences for vegetation structure. When combined, these advances hold promise for improving the scientific understanding of changes in vegetation demographics and disturbances.

Climate change and ecosystems: threats, opportunities and solutions.

The rapid anthropogenic climate change that is being experienced in the early twenty-first century is intimately entwined with the health and functioning of the biosphere. Climate change is impacting ecosystems through changes in mean conditions and in climate variability, coupled with other associated changes such as increased ocean acidification and atmospheric carbon dioxide concentrations. It also interacts with other pressures on ecosystems, including degradation, defaunation and fragmentation. There is a need to understand the ecological dynamics of these climate impacts, to identify hotspots of vulnerability and resilience and to identify management interventions that may assist biosphere resilience to climate change. At the same time, ecosystems can also assist in the mitigation of, and adaptation to, climate change. The mechanisms, potential and limits of such nature-based solutions to climate change need to be explored and quantified. This paper introduces a thematic issue dedicated to the interaction between climate change and the biosphere. It explores novel perspectives on how ecosystems respond to climate change, how ecosystem resilience can be enhanced and how ecosystems can assist in addressing the challenge of a changing climate. It draws on a Royal Society-National Academy of Sciences Forum held in Washington DC in November 2018, where these themes and issues were discussed. We conclude by identifying some priorities for academic research and practical implementation, in order to maximize the potential for maintaining a diverse, resilient and well-functioning biosphere under the challenging conditions of the twenty-first century. This article is part of the theme issue 'Climate change and ecosystems: threats, opportunities and solutions'.

Climate change, ecosystems and abrupt change: science priorities.

Ecologists have long studied patterns, directions and tempos of change, but there is a pressing need to extend current understanding to empirical observations of abrupt changes as climate warming accelerates. Abrupt changes in ecological systems (ACES)-changes that are fast in time or fast relative to their drivers-are ubiquitous and increasing in frequency. Powerful theoretical frameworks exist, yet applications in real-world landscapes to detect, explain and anticipate ACES have lagged. We highlight five insights emerging from empirical studies of ACES across diverse ecosystems: (i) ecological systems show ACES in some dimensions but not others; (ii) climate extremes may be more important than mean climate in generating ACES; (iii) interactions among multiple drivers often produce ACES; (iv) contingencies, such as ecological memory, frequency and sequence of disturbances, and spatial context are important; and (v) tipping points are often (but not always) associated with ACES. We suggest research priorities to advance understanding of ACES in the face of climate change. Progress in understanding ACES requires strong integration of scientific approaches (theory, observations, experiments and process-based models) and high-quality empirical data drawn from a diverse array of ecosystems. This article is part of the theme issue 'Climate change and ecosystems: threats, opportunities and solutions'.

Can wildland fire management alter 21st-century subalpine fire and forests in Grand Teton National Park, Wyoming, USA?

In subalpine forests of the western United States that historically experienced infrequent, high-severity fire, whether fire management can shape 21st-century fire regimes and forest dynamics to meet natural resource objectives is not known. Managed wildfire use (i.e., allowing lightning-ignited fires to burn when risk is low instead of suppressing them) is one approach for maintaining natural fire regimes and fostering mosaics of forest structure, stand age, and tree-species composition, while protecting people and property. However, little guidance exists for where and when this strategy may be effective with climate change. We simulated most of the contiguous forest in Grand Teton National Park, Wyoming, USA to ask: (1) how would subalpine fires and forest structure be different if fires had not been suppressed during the last three decades? And (2) what is the relative influence of climate change vs. fire management strategy on future fire and forests? We contrasted fire and forests from 1989 to 2098 under two fire management scenarios (managed wildfire use and fire suppression), two general circulation models (CNRM-CM5 and GFDL-ESM2M), and two representative concentration pathways (8.5 and 4.5). We found little difference between management scenarios in the number, size, or severity of fires during the last three decades. With 21st-century warming, fire activity increased rapidly, particularly after 2050, and followed nearly identical trajectories in both management scenarios. Area burned per year between 2018 and 2099 was 1,700% greater than in the last three decades (1989-2017). Large areas of forest were abruptly lost; only 65% of the original 40,178 ha of forest remained by 2098. However, forests stayed connected and fuels were abundant enough to support profound increases in burning through this century. Our results indicate that strategies emphasizing managed wildfire use, rather than suppression, will not alter climate-induced changes to fire and forests in subalpine landscapes of western North America. This suggests that managers may continue to have flexibility to strategically suppress subalpine fires without concern for long-term consequences, in distinct contrast with dry conifer forests of the Rocky Mountains and mixed conifer forest of California where maintaining low fuel loads is essential for sustaining frequent, low-severity surface fire regimes.

Comparing the effects of climate and land use on surface water quality using future watershed scenarios.

Eutrophication of freshwaters occurs in watersheds with excessive pollution of phosphorus (P). Factors that affect P cycling and transport, including climate and land use, are changing rapidly and can have legacy effects, making future freshwater quality uncertain. Focusing on the Yahara Watershed (YW) of southern Wisconsin, USA, an intensive agricultural landscape, we explored the relative influence of land use and climate on three indicators of water quality over a span of 57 years (2014-2070). The indicators included watershed-averaged P yield from the land surface, direct drainage P loads to a lake, and average summertime lake P concentration. Using biophysical model simulations of future watershed scenarios, we found that climate exerted a stronger influence than land use on all three indicators, yet land use had an important role in influencing long term outcomes for each. Variations in P yield due to land use exceeded those due to climate in 36 of 57 years, whereas variations in load and lake total P concentration due to climate exceeded those due to land use in 54 of 57 years, and 52 of 57 years, respectively. The effect of land use was thus strongest for P yield off the landscape and attenuated in the stream and lake aquatic systems where the influence of weather variability was greater. Overall these findings underscore the dominant role of climate in driving inter-annual nutrient fluxes within the hydrologic network and suggest a challenge for land use to influence water quality within streams and lakes over timescales less than a decade. Over longer timescales, reducing applications of P throughout the watershed was an effective management strategy under all four climates investigated, even during decades with wetter conditions and more frequent extreme precipitation events.

Short-interval severe fire erodes the resilience of subalpine lodgepole pine forests.

Subalpine forests in the northern Rocky Mountains have been resilient to stand-replacing fires that historically burned at 100- to 300-year intervals. Fire intervals are projected to decline drastically as climate warms, and forests that reburn before recovering from previous fire may lose their ability to rebound. We studied recent fires in Greater Yellowstone (Wyoming, United States) and asked whether short-interval (<30 years) stand-replacing fires can erode lodgepole pine (Pinus contorta var. latifolia) forest resilience via increased burn severity, reduced early postfire tree regeneration, reduced carbon stocks, and slower carbon recovery. During 2016, fires reburned young lodgepole pine forests that regenerated after wildfires in 1988 and 2000. During 2017, we sampled 0.25-ha plots in stand-replacing reburns (n = 18) and nearby young forests that did not reburn (n = 9). We also simulated stand development with and without reburns to assess carbon recovery trajectories. Nearly all prefire biomass was combusted ("crown fire plus") in some reburns in which prefire trees were dense and small (≤4-cm basal diameter). Postfire tree seedling density was reduced sixfold relative to the previous (long-interval) fire, and high-density stands (>40,000 stems ha-1) were converted to sparse stands (<1,000 stems ha-1). In reburns, coarse wood biomass and aboveground carbon stocks were reduced by 65 and 62%, respectively, relative to areas that did not reburn. Increased carbon loss plus sparse tree regeneration delayed simulated carbon recovery by >150 years. Forests did not transition to nonforest, but extreme burn severity and reduced tree recovery foreshadow an erosion of forest resilience.

Scale-dependent interactions between tree canopy cover and impervious surfaces reduce daytime urban heat during summer.

As cities warm and the need for climate adaptation strategies increases, a more detailed understanding of the cooling effects of land cover across a continuum of spatial scales will be necessary to guide management decisions. We asked how tree canopy cover and impervious surface cover interact to influence daytime and nighttime summer air temperature, and how effects vary with the spatial scale at which land-cover data are analyzed (10-, 30-, 60-, and 90-m radii). A bicycle-mounted measurement system was used to sample air temperature every 5 m along 10 transects (∼7 km length, sampled 3-12 times each) spanning a range of impervious and tree canopy cover (0-100%, each) in a midsized city in the Upper Midwest United States. Variability in daytime air temperature within the urban landscape averaged 3.5 °C (range, 1.1-5.7 °C). Temperature decreased nonlinearly with increasing canopy cover, with the greatest cooling when canopy cover exceeded 40%. The magnitude of daytime cooling also increased with spatial scale and was greatest at the size of a typical city block (60-90 m). Daytime air temperature increased linearly with increasing impervious cover, but the magnitude of warming was less than the cooling associated with increased canopy cover. Variation in nighttime air temperature averaged 2.1 °C (range, 1.2-3.0 °C), and temperature increased with impervious surface. Effects of canopy were limited at night; thus, reduction of impervious surfaces remains critical for reducing nighttime urban heat. Results suggest strategies for managing urban land-cover patterns to enhance resilience of cities to climate warming.

Feast not famine: Nitrogen pools recover rapidly in 25-yr-old postfire lodgepole pine.

The extent of young postfire conifer forests is growing throughout western North America as the frequency and size of high-severity fires increase, making it important to understand ecosystem structure and function in early seral forests. Understanding nitrogen (N) dynamics during postfire stand development is especially important because northern conifers are often N limited. We resampled lodgepole pine (Pinus contorta var. latifolia) stands that regenerated naturally after the 1988 fires in Yellowstone National Park (Wyoming, USA) to ask (1) How have N pools and fluxes changed over a decade (15 to 25 yr postfire) of very rapid forest growth? (2) At postfire year 25, how do N pools and fluxes vary with lodgepole pine density and productivity? Lodgepole pine foliage, litter (annual litterfall, forest-floor litter), and mineral soils were sampled in 14 plots (0.25 ha) that varied in postfire lodgepole pine density (1,500 to 344,000 stems/ha) and aboveground net primary production (ANPP; 1.4 to 16.1 Mg·ha-1 ·yr-1 ). Counter to expectation, foliar N concentrations in lodgepole pine current-year and composite needles (1.33 and 1.11% N, respectively) had not changed over time. Further, all measured ecosystem N pools increased substantially: foliar N increased to 89 kg N/ha (+93%), O-horizon N increased to 39 kg N/ha (+38%), and mineral soil percent total N (0-15 cm) increased to 0.08% (+33%). Inorganic N availability also increased to 0.69 µg N·[g resin]-1 ·d-1 (+165%). Thus, soil N did not decline as live biomass N pools increased. Among stands, biomass N pools at postfire year 25 remained strongly influenced by early postfire tree density: foliar and litterfall N concentrations declined with lodgepole pine density and ANPP, but the foliar N pool increased. Lodgepole pine ANPP correlated negatively with annual resin-sorbed N, and we found no indication of widespread N limitation. The large increases in N pools cannot be explained by atmospheric N deposition or presence of known N fixers. These results suggest an unmeasured N source and are consistent with recent reports of N fixation in young lodgepole pine.

Effects of bird community dynamics on the seasonal distribution of cultural ecosystem services.

Biodiversity-based cultural ecosystem services (CES), such as birdwatching, are strongly influenced by biotic community dynamics. However, CES models are largely static, relying on single estimates of species richness or land-use/land-cover proxies, and may be inadequate for landscape management of CES supply. Using bird survey data from the Appalachian Mountains (USA), we developed spatial-temporal models of five CES indicators (total bird species richness, and richness of migratory, infrequent, synanthrope, and resident species), reflecting variation in birdwatcher preferences. We analyzed seasonal shifts in birdwatching supply and how those shifts impacted public access to projected birdwatching hotspots. Landscape patterns of CES supply differed substantially among indicators, leading to opposing conclusions about locations of highest birdwatching supply. Total species richness hotspots seldom overlapped with hotspots of migratory or infrequent species. Public access to CES hotspots varied seasonally. Our study suggests that simple, static biodiversity metrics may overlook spatial dynamics important to CES users.