High-resolution soil sampling reveals the pattern of biological weathering and soil formation under trees.

Trees contribute to bedrock weathering in a variety of ways. However, evaluating their full impact is complicated by a lack of direct observation of unexposed root systems of individual trees, especially when the scale of the analysis goes down to the level of microbiomes. In the present study, we investigated the contribution of tree root systems to bioweathering and soil production at the macro- and microscale. Soil profiles developed under trees on granite bedrock were investigated in two parts of the Sudety Mountains, SW Poland: the Rudawy Janowickie Mountains, and the Stolowe Mountains. Soil profiles were gradually excavated and soil samples collected from pre-defined positions of the root zone: 1) bulk soil, 2) rhizosphere, 3) cracks, 4) topsoil, and 5) control positions. In total, we analyzed 103 samples for soil chemistry and microbiological activity. In addition, we analyzed 19 samples using XRF (X-ray Fluorescence). Four parent rock samples, in the form of thin-sections, were the subject of mineralogical evaluation. Soil analyses included: total organic carbon (C) and nitrogen (N) content, soil pHH2O, soluble iron (Fed), and aluminum (Ald), non-crystalline (amorphous) iron (Feox), and aluminum (Alox). For microbiological analyses, we used a Biolog (EcoPlate) system to determine the functional diversity of soil microorganisms. We evaluated the results on soil chemistry and microbiological activity statistically by principal component analysis (PCA) and redundancy analysis (RDA). Differences between soil sampling positions were assessed using a non-parametric Kruskal-Wallis (K-W) rank sum test and a post-hoc pairwise Dunn test. Trees developed different root architectures, likely shaped by the depth to bedrock and its pre-existing net of fractures and fissures. Tree roots were able to enter bedrock cracks at one study site (at Pstrazna, Stolowe Mountains). The soil profile was too deep for root system penetration at the second study site (Mt Janska, Rudawy Janowickie Mountains, RJM). The rhizospheric soil along the roots had significantly different chemical properties compared to non-rhizospheric soil types. At Mt. Janska, soil differed from the crack soil in terms of Alox (pHolm-adj. < 0.0006) and Feox (pHolm-adj. < 0.004), and from the bulk soil (pHolm-adj. < 0.02) and topsoil (pHolm-adj. < 0.007). In addition, at Pstrazna, the soil differed from the control soil in terms of C (pHolm-adj. < 0.009) and soil pHH2O (pHolm-adj. < 0.0008) and from the topsoil in terms of soil pHH2O. The highest metabolic activity was in cracks at Mt. Janska and in control samples from Pstrazna. In general, the spatial distribution of soil microbial activity, and the weathering that results from that portion of the soil biome, is spatially heterogeneous and appears to be partially determined by the interaction of root growth and bedrock fracture patterns.

Disturbance theory for ecosystem ecologists: A primer.

Understanding what regulates ecosystem functional responses to disturbance is essential in this era of global change. However, many pioneering and still influential disturbance-related theorie proposed by ecosystem ecologists were developed prior to rapid global change, and before tools and metrics were available to test them. In light of new knowledge and conceptual advances across biological disciplines, we present four disturbance ecology concepts that are particularly relevant to ecosystem ecologists new to the field: (a) the directionality of ecosystem functional response to disturbance; (b) functional thresholds; (c) disturbance-succession interactions; and (d) diversity-functional stability relationships. We discuss how knowledge, theory, and terminology developed by several biological disciplines, when integrated, can enhance how ecosystem ecologists analyze and interpret functional responses to disturbance. For example, when interpreting thresholds and disturbance-succession interactions, ecosystem ecologists should consider concurrent biotic regime change, non-linearity, and multiple response pathways, typically the theoretical and analytical domain of population and community ecologists. Similarly, the interpretation of ecosystem functional responses to disturbance requires analytical approaches that recognize disturbance can promote, inhibit, or fundamentally change ecosystem functions. We suggest that truly integrative approaches and knowledge are essential to advancing ecosystem functional responses to disturbance.

Weathering and soil production under trees growing on sandstones - The role of tree roots in soil formation.

Rock weathering drives both landform formation and soil production/evolution. The less studied biological component of weathering and soil production caused by tree root systems is the main focus of the present study. Weathering by trees, which likely has been important in soil formation since the first trees emerged in the middle and late Devonian, is accomplished through both physical and biological means, like acids excreted by plants and exudates from associated bacterial communities. However, these processes are relatively poorly known. We assessed the impact of tree roots and associated microbiota on the potential level of biological weathering. Three research plots were selected in two sandstone regions in Poland. Two plots were in the Stolowe Mountains (Zlotno, Batorów), a tableland built of Cretaceous sandstones. The third plot (Zegiestów) was in the Sacz Beskidy Mountains, the Carpathians. Soil samples were taken from tree root zones of Norway spruces from predefined sampling positions. Soils from non-tree control positions were also sampled. Soil samples were a subject of laboratory analyses which included the content of Fe and Al (amorphous and labile forms), carbon (C), nitrogen (N), and soil pH. The microbial functional diversity of soil microorganisms was determined using the Biolog (EcoPlate) system. Rock fragments were collected for mineralogical and a subject of optical microscopy and cathodoluminescence analyses in order to examine their mineralogical composition. Significant differences (pHolm-corrected < 0.05) between sample locations were found mostly for the Zegiestów plot: Soils at control positions differed from the crack and bulk soil sample positions in terms of C, N, C/N, and pH. Tree roots were able to develop a great variety of sizes and forms by following the existing net of bedrock discontinuities and hillslope microrelief. They developed along the most accessible surfaces, and caused rockcliff retreat and scree slope formation. These two features can be considered as initial stages of soil production. Trees add to the complexity of the soil system and allow formation of rhizospheric soils, and horizons rich in organic matter which are zones of a high microbial activity. However, as our study shows, rock cracks with roots cannot be considered as zones of microbial weathering. In addition, C content and microbial activity decreases with depth but can stay on a high level along living and dead roots. When entering rock fractures, they change the intensity of biomechanical weathering and soil properties. The highest biological activity of microorganisms was found in the control samples. Overall, tree roots do change the pattern of soil formation and explain the existing pattern of soil chemical properties, microbial activity, and potentially biological weathering intensity, and the intensity of those processes in correlation with root presence varies in space.

Reduced fire severity offers near-term buffer to climate-driven declines in conifer resilience across the western United States.

Increasing fire severity and warmer, drier postfire conditions are making forests in the western United States (West) vulnerable to ecological transformation. Yet, the relative importance of and interactions between these drivers of forest change remain unresolved, particularly over upcoming decades. Here, we assess how the interactive impacts of changing climate and wildfire activity influenced conifer regeneration after 334 wildfires, using a dataset of postfire conifer regeneration from 10,230 field plots. Our findings highlight declining regeneration capacity across the West over the past four decades for the eight dominant conifer species studied. Postfire regeneration is sensitive to high-severity fire, which limits seed availability, and postfire climate, which influences seedling establishment. In the near-term, projected differences in recruitment probability between low- and high-severity fire scenarios were larger than projected climate change impacts for most species, suggesting that reductions in fire severity, and resultant impacts on seed availability, could partially offset expected climate-driven declines in postfire regeneration. Across 40 to 42% of the study area, we project postfire conifer regeneration to be likely following low-severity but not high-severity fire under future climate scenarios (2031 to 2050). However, increasingly warm, dry climate conditions are projected to eventually outweigh the influence of fire severity and seed availability. The percent of the study area considered unlikely to experience conifer regeneration, regardless of fire severity, increased from 5% in 1981 to 2000 to 26 to 31% by mid-century, highlighting a limited time window over which management actions that reduce fire severity may effectively support postfire conifer regeneration.

Reimagine fire science for the anthropocene.

Fire is an integral component of ecosystems globally and a tool that humans have harnessed for millennia. Altered fire regimes are a fundamental cause and consequence of global change, impacting people and the biophysical systems on which they depend. As part of the newly emerging Anthropocene, marked by human-caused climate change and radical changes to ecosystems, fire danger is increasing, and fires are having increasingly devastating impacts on human health, infrastructure, and ecosystem services. Increasing fire danger is a vexing problem that requires deep transdisciplinary, trans-sector, and inclusive partnerships to address. Here, we outline barriers and opportunities in the next generation of fire science and provide guidance for investment in future research. We synthesize insights needed to better address the long-standing challenges of innovation across disciplines to (i) promote coordinated research efforts; (ii) embrace different ways of knowing and knowledge generation; (iii) promote exploration of fundamental science; (iv) capitalize on the "firehose" of data for societal benefit; and (v) integrate human and natural systems into models across multiple scales. Fire science is thus at a critical transitional moment. We need to shift from observation and modeled representations of varying components of climate, people, vegetation, and fire to more integrative and predictive approaches that support pathways toward mitigating and adapting to our increasingly flammable world, including the utilization of fire for human safety and benefit. Only through overcoming institutional silos and accessing knowledge across diverse communities can we effectively undertake research that improves outcomes in our more fiery future.

From canopy to seed: Loss of snow drives directional changes in forest composition.

Climate change is altering the conditions for tree recruitment, growth, and survival, and impacting forest community composition. Across southeast Alaska, USA, and British Columbia, Canada, Callitropsis nootkatensis (Alaska yellow-cedar) is experiencing extensive climate change-induced canopy mortality due to fine-root death during soil freezing events following warmer winters and the loss of insulating snowpack. Here, we examine the effects of ongoing, climate-driven canopy mortality on forest community composition and identify potential shifts in stand trajectories due to the loss of a single canopy species. We sampled canopy and regenerating forest communities across the extent of C. nootkatensis decline in southeast Alaska to quantify the effects of climate, community, and stand-level drivers on C. nootkatensis canopy mortality and regeneration as well as postdecline regenerating community composition. Across the plot network, C. nootkatensis exhibited significantly higher mortality than co-occurring conifers across all size classes and locations. Regenerating community composition was highly variable but closely related to the severity of C. nootkatensis mortality. Callitropsis nootkatensis canopy mortality was correlated with winter temperatures and precipitation as well as local soil drainage, with regenerating community composition and C. nootkatensis regeneration abundances best explained by available seed source. In areas of high C. nootkatensis mortality, C. nootkatensis regeneration was low and replaced by Tsuga. Our study suggests that climate-induced forest mortality is driving alternate successional pathways in forests where C. nootkatensis was once a major component. These pathways are likely to lead to long-term shifts in forest community composition and stand dynamics. Our analysis fills a critical knowledge gap on forest ecosystem response and rearrangement following the climate-driven decline of a single species, providing new insight into stand dynamics in a changing climate. As tree species across the globe are increasingly stressed by climate change-induced alteration of suitable habitat, identifying the autecological factors contributing to successful regeneration, or lack thereof, will provide key insight into forest resilience and persistence on the landscape.

Integrating Subjective and Objective Dimensions of Resilience in Fire-Prone Landscapes.

Resilience has become a common goal for science-based natural resource management, particularly in the context of changing climate and disturbance regimes. Integrating varying perspectives and definitions of resilience is a complex and often unrecognized challenge to applying resilience concepts to social-ecological systems (SESs) management. Using wildfire as an example, we develop a framework to expose and separate two important dimensions of resilience: the inherent properties that maintain structure, function, or states of an SES and the human perceptions of desirable or valued components of an SES. In doing so, the framework distinguishes between value-free and human-derived, value-explicit dimensions of resilience. Four archetypal scenarios highlight that ecological resilience and human values do not always align and that recognizing and anticipating potential misalignment is critical for developing effective management goals. Our framework clarifies existing resilience theory, connects literature across disciplines, and facilitates use of the resilience concept in research and land-management applications.

Long-term exposure to more frequent disturbances increases baseline carbon in some ecosystems: Mapping and quantifying the disturbance frequency-ecosystem C relationship.

Disturbance regimes have a major influence on the baseline carbon that characterizes any particular ecosystem. Often regimes result in lower average regional baseline C (compared to those same systems if the disturbance processes were lessened/removed). However, in infrequently disturbed systems the role of disturbance as a "background" process that influences broad-scale, baseline C levels is often neglected. Long-term chronosequences suggest disturbances in these systems may serve to increase regional biomass C stocks by maintaining productivity. However, that inference has not been tested spatially. Here, the large forested system of southeast Alaska, USA, is utilized to 1) estimate baseline regional C stocks, 2) test the fundamental disturbance-ecosystem C relationship, 3) estimate the cumulative impact of disturbances on baseline C. Using 1491 ground points with carbon measurements and a novel way of mapping disturbance regimes, the relationship between total biomass C, disturbance exposure, and climate was analyzed statistically. A spatial model was created to determine regional C and compare different disturbance scenarios. In this infrequently disturbed ecosystem, higher disturbance exposure is correlated with higher biomass C, supporting the hypothesis that disturbances maintain productivity at broad scales. The region is estimated to potentially contain a baseline 1.21-1.52 Pg biomass C (when unmanaged). Removal of wind and landslides from the model resulted in lower net C stocks (-2 to -19% reduction), though the effect was heterogeneous on finer scales. There removal of landslides alone had a larger effect then landslide and wind combined removal. The relationship between higher disturbance exposure and higher biomass within the broad ecosystem (which, on average, has a very low disturbance frequency) suggest that disturbances can serve maintain higher levels of productivity in infrequently disturbed but very C dense ecosystems. Carbon research in other systems, especially those where disturbances are infrequent relative to successional processes, should consider the role of disturbances in maintaining baseline ecosystem productivity.

Patterns and drivers of recent disturbances across the temperate forest biome.

Increasing evidence indicates that forest disturbances are changing in response to global change, yet local variability in disturbance remains high. We quantified this considerable variability and analyzed whether recent disturbance episodes around the globe were consistently driven by climate, and if human influence modulates patterns of forest disturbance. We combined remote sensing data on recent (2001-2014) disturbances with in-depth local information for 50 protected landscapes and their surroundings across the temperate biome. Disturbance patterns are highly variable, and shaped by variation in disturbance agents and traits of prevailing tree species. However, high disturbance activity is consistently linked to warmer and drier than average conditions across the globe. Disturbances in protected areas are smaller and more complex in shape compared to their surroundings affected by human land use. This signal disappears in areas with high recent natural disturbance activity, underlining the potential of climate-mediated disturbance to transform forest landscapes.

Determining the size of a complete disturbance landscape: multi-scale, continental analysis of forest change.

The scale of investigation for disturbance-influenced processes plays a critical role in theoretical assumptions about stability, variance, and equilibrium, as well as conservation reserve and long-term monitoring program design. Critical consideration of scale is required for robust planning designs, especially when anticipating future disturbances whose exact locations are unknown. This research quantified disturbance proportion and pattern (as contagion) at multiple scales across North America. This pattern of scale-associated variability can guide selection of study and management extents, for example, to minimize variance (measured as standard deviation) between any landscapes within an ecoregion. We identified the proportion and pattern of forest disturbance (30 m grain size) across multiple landscape extents up to 180 km2. We explored the variance in proportion of disturbed area and the pattern of that disturbance between landscapes (within an ecoregion) as a function of the landscape extent. In many ecoregions, variance between landscapes within an ecoregion was minimal at broad landscape extents (low standard deviation). Gap-dominated regions showed the least variance, while fire-dominated showed the largest. Intensively managed ecoregions displayed unique patterns. A majority of the ecoregions showed low variance between landscapes at some scale, indicating an appropriate extent for incorporating natural regimes and unknown future disturbances was identified. The quantification of the scales of disturbance at the ecoregion level provides guidance for individuals interested in anticipating future disturbances which will occur in unknown spatial locations. Information on the extents required to incorporate disturbance patterns into planning is crucial for that process.

A foundation of ecology rediscovered: 100 years of succession on the William S. Cooper plots in Glacier Bay, Alaska.

Understanding plant community succession is one of the original pursuits of ecology, forming some of the earliest theoretical frameworks in the field. Much of this was built on the long-term research of William S. Cooper, who established a permanent plot network in Glacier Bay, Alaska, in 1916. This study now represents the longest-running primary succession plot network in the world. Permanent plots are useful for their ability to follow mechanistic change through time without assumptions inherent in space-for-time (chronosequence) designs. After 100-yr, these plots show surprising variety in species composition, soil characteristics (carbon, nitrogen, depth), and percent cover, attributable to variation in initial vegetation establishment first noted by Cooper in the 1916-1923 time period, partially driven by dispersal limitations. There has been almost a complete community composition replacement over the century and general species richness increase, but the effective number of species has declined significantly due to dominance of Salix species which established 100-yr prior (the only remaining species from the original cohort). Where Salix dominates, there is no establishment of "later" successional species like Picea. Plots nearer the entrance to Glacier Bay, and thus closer to potential seed sources after the most recent glaciation, have had consistently higher species richness for 100 yr. Age of plots is the best predictor of soil N content and C:N ratio, though plots still dominated by Salix had lower overall N; soil accumulation was more associated with dominant species. This highlights the importance of contingency and dispersal in community development. The 100-yr record of these plots, including species composition, spatial relationships, cover, and observed interactions between species provides a powerful view of long-term primary succession.

Emerging climate-driven disturbance processes: widespread mortality associated with snow-to-rain transitions across 10° of latitude and half the range of a climate-threatened conifer.

Climate change is causing rapid changes to forest disturbance regimes worldwide. While the consequences of climate change for existing disturbance processes, like fires, are relatively well studied, emerging drivers of disturbance such as snow loss and subsequent mortality are much less documented. As the climate warms, a transition from winter snow to rain in high latitudes will cause significant changes in environmental conditions such as soil temperatures, historically buffered by snow cover. The Pacific coast of North America is an excellent test case, as mean winter temperatures are currently at the snow-rain threshold and have been warming for approximately 100 years post-Little Ice Age. Increased mortality in a widespread tree species in the region has been linked to warmer winters and snow loss. Here, we present the first high-resolution range map of this climate-sensitive species, Callitropsis nootkatensis (yellow-cedar), and document the magnitude and location of observed mortality across Canada and the United States. Snow cover loss related mortality spans approximately 10° latitude (half the native range of the species) and 7% of the overall species range and appears linked to this snow-rain transition across its range. Mortality is commonly >70% of basal area in affected areas, and more common where mean winter temperatures is at or above the snow-rain threshold (>0 °C mean winter temperature). Approximately 50% of areas with a currently suitable climate for the species (<-2 °C) are expected to warm beyond that threshold by the late 21st century. Regardless of climate change scenario, little of the range which is expected to remain suitable in the future (e.g., a climatic refugia) is in currently protected landscapes (<1-9%). These results are the first documentation of this type of emerging climate disturbance and highlight the difficulties of anticipating novel disturbance processes when planning for conservation and management.

Spatial and topographic trends in forest expansion and biomass change, from regional to local scales.

Natural forest growth and expansion are important carbon sequestration processes globally. Climate change is likely to increase forest growth in some regions via CO2 fertilization, increased temperatures, and altered precipitation; however, altered disturbance regimes and climate stress (e.g. drought) will act to reduce carbon stocks in forests as well. Observations of asynchrony in forest change is useful in determining current trends in forest carbon stocks, both in terms of forest density (e.g. Mg ha(-1) ) and spatially (extent and location). Monitoring change in natural (unmanaged) areas is particularly useful, as while afforestation and recovery from historic land use are currently large carbon sinks, the long-term viability of those sinks depends on climate change and disturbance dynamics at their particular location. We utilize a large, unmanaged biome (>135 000 km(2) ) which spans a broad latitudinal gradient to explore how variation in location affects forest density and spatial patterning: the forests of the North American temperate rainforests in Alaska, which store >2.8 Pg C in biomass and soil, equivalent to >8% of the C in contiguous US forests. We demonstrate that the regional biome is shifting; gains exceed losses and are located in different spatio-topographic contexts. Forest gains are concentrated on northerly aspects, lower elevations, and higher latitudes, especially in sheltered areas, whereas loss is skewed toward southerly aspects and lower latitudes. Repeat plot-scale biomass data (n = 759) indicate that within-forest biomass gains outpace losses (live trees >12.7 cm diameter, 986 Gg yr(-1) ) on gentler slopes and in higher latitudes. This work demonstrates that while temperate rainforest dynamics occur at fine spatial scales (<1000 m(2) ), the net result of thousands of individual events is regionally patterned change. Correlations between the disturbance/establishment imbalance and biomass accumulation suggest the potential for relatively rapid biome shifts and biomass changes.

Evaluating the utility and seasonality of NDVI values for assessing post-disturbance recovery in a subalpine forest.

Forest disturbances around the world have the potential to alter forest type and cover, with impacts on diversity, carbon storage, and landscape composition. These disturbances, especially fire, are common and often large, making ground investigation of forest recovery difficult. Remote sensing offers a means to monitor forest recovery in real time, over the entire landscape. Typically, recovery monitoring via remote sensing consists of measuring vegetation indices (e.g., NDVI) or index-derived metrics, with the assumption that recovery in NDVI (for example) is a meaningful measure of ecosystem recovery. This study tests that assumption using MODIS 16-day imagery from 2000 to 2010 in the area of the Colorado's Routt National Forest Hinman burn (2002) and seedling density counts taken in the same area. Results indicate that NDVI is rarely correlated with forest recovery, and is dominated by annual and perennial forb cover, although topography complicates analysis. Utility of NDVI as a means to delineate areas of recovery or non-recovery are in doubt, as bootstrapped analysis indicates distinguishing power only slightly better than random. NDVI in revegetation analyses should carefully consider the ecology and seasonal patterns of the system in question.