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DNA metabarcoding and video camera collars yield different inferences about the summer diet of an arctic ungulate

The diets of wild ungulates are a foundational component of their ecology, influencing their behavior, body condition, and demography. With changing environmental conditions, there is a significant need to identify important forage items for ungulates, but this has often proved challenging. Declines in several barren-ground caribou herds across the North American Arctic have raised concerns about the influence of climate change on caribou forage conditions. Shifts in plant phenology, biomass, quality, and composition may be influencing caribou diets and subsequently affecting their body condition and demographic rates. Although forage is a primary driver of barren-ground caribou behavior and population dynamics, there is limited recent information about the specific foods they consume, and uncertainty about appropriate methods for identifying those foods. Investigators are increasingly using fecal DNA metabarcoding and video camera collars to assess ungulate diets, but comparative studies of these approaches are lacking. To examine the summer diets of barren-ground caribou, we used both fecal metabarcoding and video camera collars to identify forage used by the Porcupine caribou herd, which spans the Alaska–Yukon border. In 2021, we sampled the diets of adult females by collecting fecal samples and observing collar videos during 4 sampling occasions. We found that caribou consumed very specific forage items, and those items varied markedly across the growing season. Caribou predominantly consumed graminoids and lichens during early summer, and shrubs and forbs later in the season. Metabarcoding and video data provided significantly different estimates of diet for all taxonomic levels we evaluated, and inferences from the two approaches were often disparate. Metabarcoding failed to detect some items frequently consumed in videos, such as lichens, and indicated high use of other items rarely consumed, such as mosses. We found that video data provided greater taxonomic diversity and resolution for vascular plants and lichens, and more closely aligned with past research and biological expectations than data from fecal metabarcoding. Additional research is needed to be able to use these methods to identify the biomass of different forage items consumed.

Alaska, Yukon↗

Improving ecological restoration to curb biotic invasion - A practical guide

Common practices for invasive species control and management include physical, chemical, and biological approaches. The first two approaches have clear limitations and may lead to unintended (negative) consequences, unless carefully planned and implemented. For example, physical removal rarely completely eradicates the targeted invasive species and can cause disturbances that facilitate new invasions by nonnative species from nearby habitats. Chemical treatments can harm native, and especially rare, species through unanticipated side effects. Biological methods may be classified as biocontrol and the ecological approach. Similar to physical and chemical methods, biocontrol also has limitations and sometimes leads to unintended consequences. Therefore, a relatively safer and more practical choice may be the ecological approach, which has two major components: (1) restoration of native species and (2) biomass manipulation of the restored community, such as selective grazing or prescribed burning (to achieve and maintain viable population sizes). Restoration requires well-planned and implemented planting designs that consider alpha-, beta-, and gamma-diversity and the abundance of native and invasive component species at local, landscape, and regional levels. Given the extensive destruction or degradation of natural habitats around the world, restoration could be most effective for enhancing ecosystem resilience and resistance to biotic invasions. At the same time, ecosystems in human-dominated landscapes, especially those newly restored, require close monitoring and careful intervention (e.g., through biomass manipulation), especially when successional trajectories are not moving as intended. Biomass management frequently uses prescribed burning, grazing, harvesting, and thinning to maintain overall ecosystem health and sustainability. Thus, the resulting optimal, balanced, and relatively stable ecological conditions could more effectively limit the spread and establishment of invasive species. Here we review the literature (especially within the last decade) on ecological approaches that involve biodiversity, biomass, and productivity, three key community/ecosystem variables that reciprocally influence one another. We focus on the common and most feasible ecological practices that can aid in resisting new invasions and/or suppressing the dominance of existing invasive species. We contend that, because of the strong influences from neighboring areas (i.e., as exotic species pools), local restoration and management efforts in the future need to consider the regional context and projected climate changes.

Invasive Plant Science and Management↗

Beyond just sea-level rise: Considering macroclimatic drivers within coastal wetland vulnerability assessments to climate change

Due to their position at the land-sea interface, coastal wetlands are vulnerable to many aspects of climate change. However, climate change vulnerability assessments for coastal wetlands generally focus solely on sea-level rise without considering the effects of other facets of climate change. Across the globe and in all ecosystems, macroclimatic drivers (e.g., temperature and rainfall regimes) greatly influence ecosystem structure and function. Macroclimatic drivers have been the focus of climate-change related threat evaluations for terrestrial ecosystems, but largely ignored for coastal wetlands. In some coastal wetlands, changing macroclimatic conditions are expected to result in foundation plant species replacement, which would affect the supply of certain ecosystem goods and services and could affect ecosystem resilience. As examples, we highlight several ecological transition zones where small changes in macroclimatic conditions would result in comparatively large changes in coastal wetland ecosystem structure and function. Our intent in this communication is not to minimize the importance of sea-level rise. Rather, our overarching aim is to illustrate the need to also consider macroclimatic drivers within vulnerability assessments for coastal wetlands.

Global Change Biology↗

Vulnerability of riparian ecosystems to elevated CO 2 and climate change in arid and semiarid western North America

Riparian ecosystems, already greatly altered by water management, land development, and biological invasion, are being further altered by increasing atmospheric CO 2 concentrations ([CO 2 ]) and climate change, particularly in arid and semiarid (dryland) regions. In this literature review, we (1) summarize expected changes in [CO 2 ], climate, hydrology, and water management in dryland western North America, (2) consider likely effects of those changes on riparian ecosystems, and (3) identify critical knowledge gaps. Temperatures in the region are rising and droughts are becoming more frequent and intense. Warmer temperatures in turn are altering river hydrology: advancing the timing of spring snow melt floods, altering flood magnitudes, and reducing summer and base flows. Direct effects of increased [CO 2 ] and climate change on riparian ecosystems may be similar to effects in uplands, including increased heat and water stress, altered phenology and species geographic distributions, and disrupted trophic and symbiotic interactions. Indirect effects due to climate-driven changes in streamflow, however, may exacerbate the direct effects of warming and increase the relative importance of moisture and fluvial disturbance as drivers of riparian ecosystem response to global change. Together, climate change and climate-driven changes in streamflow are likely to reduce abundance of dominant, native, early-successional tree species, favor herbaceous species and both drought-tolerant and late-successional woody species (including many introduced species), reduce habitat quality for many riparian animals, and slow litter decomposition and nutrient cycling. Climate-driven changes in human water demand and associated water management may intensify these effects. On some regulated rivers, however, reservoir releases could be managed to protect riparian ecosystem. Immediate research priorities include determining riparian species' environmental requirements and monitoring riparian ecosystems to allow rapid detection and response to undesirable ecological change.

Global Change Biology↗

U.S. Geological Survey science vision for native freshwater mussel research in the United States

Executive Summary North America is a global center for native freshwater mussel (order Unionida, hereinafter “mussels”) diversity, with more than 350 species. Mussels are among the most imperiled fauna on the planet. Reasons for both local and widespread declines in mussels are mostly unknown, although the threats may include habitat loss and fragmentation, diseases, environmental contaminants, altered flow regimes, migration barriers to larval hosts, non-native species, and climate change. Over the past three decades, research on mussels has been substantial. Nevertheless, current conservation and management efforts are limited by significant information gaps. For example, the effects of emerging stressors on mussels are largely unknown and identifying when habitats are rehabilitated and suitable for reestablishment of mussels remains challenging. Additionally, historical and current information on the distribution, taxonomy, and life histories are often unreliable or lacking altogether, and more reliable information is needed for many species. We identified focal research themes, goals, and objectives where research on mussels is needed based on information gaps identified through conversations with resource partners across local, regional, and national organizations. Research on biodiversity seeks to enhance the diversity of mussel species and populations to support healthy aquatic ecosystems. Research on emerging stressors seeks to improve the understanding of how mussel species, populations, and communities respond to emerging stressors, including environmental contaminants and climate change. Research on conservation seeks to enhance the recovery of species and populations and to identify data gaps limiting the conservation of mussels and their habitats. Mussels are in urgent need of proactive conservation because they are an integral part of our natural heritage, enhance biodiversity, and provide vital ecological services that support freshwater ecosystems. The U.S. Geological Survey (USGS) has been, and continues to be, a leader in mussel research. Although the USGS is well suited to address the broad-scale multidisciplinary research needed to conserve mussels, the USGS has had substantial loss of scientists with mussel expertise over the past 20 years. However, the breadth of the USGS expertise on mussels can be leveraged internally across other USGS mission and program areas and externally across research partners. Given the breadth and scope of the issues facing mussels across the United States, the research themes outlined in this science vision can only be accomplished through extensive collaborations between the USGS and the full spectrum of natural resource partners, including other Federal and State agencies, Tribal organizations, universities, industries, and nongovernmental organizations.

Circular↗

High throughput computing: a solution for scientific analysis

Public land management agencies continually face resource management problems that are exacerbated by climate warming, land-use change, and other human activities. As the U.S. Geological Survey (USGS) Fort Collins Science Center (FORT) works with managers in U.S. Department of the Interior (DOI) agencies and other federal, state, and private entities, researchers are finding that the science needed to address these complex ecological questions across time and space produces substantial amounts of data. The additional data and the volume of computations needed to analyze it require expanded computing resources well beyond single- or even multiple-computer workstations. To meet this need for greater computational capacity, FORT investigated how to resolve the many computational shortfalls previously encountered when analyzing data for such projects. Our objectives included finding a solution that would: harness existing Computer Processing Units (CPUs) when they're idle to run multiple jobs concurrently, which reduces the overall processing time without requiring additional hardware; offer an effective, centralized job-management system; handle job failures due to hardware, software, or network interruptions (obviating the need to manually resubmit the job after each stoppage); be affordable; and most importantly, allow us to complete very large, complex analyses that otherwise would not even be possible. In short, we envisioned a job-management system that would take advantage of unused FORT CPUs within a local area network (LAN) to effectively distribute and run highly complex analytical processes. What we found was a solution that uses High Throughput Computing (HTC) and High Performance Computing (HPC) systems to do exactly that (Figure 1).

Report↗

WICCI Wildlife Working Group Report

Wisconsin is world-renowned for its diversity of ecological landscapes and wildlife populations. The northern forests, southern prairies, and interior and coastal wetlands of the state are home to more than 500 terrestrial animal species. These animals supply the Wisconsin public with aesthetic, cultural, and economic benefits; our identity and economy are intertwined with these natural resources. Climate change is altering the behavior, distribution, development, reproduction, and survival of these animal populations. In turn, these changes will alter the aesthetic, cultural, and economic benefits we receive from them. The focus of the Wildlife Working Group is to document past and current impacts, anticipate changes in wildlife distribution and abundance, and develop adaptation strategies to maintain the vitality and diversity of Wisconsin's wildlife populations.

Report↗

Alternative stable states in inherently unstable systems

Alternative stable states are nontransitory states within which communities can exist. However, even highly dynamic communities can be viewed within the framework of stable‐state theory if an appropriate “ecologically relevant” time scale is identified. The ecologically relevant time scale for dynamic systems needs to conform to the amount of time needed for a system's community to complete an entire cycle through its normal range of variation. For some systems, the ecologically relevant period can be relatively short (eg, tidal systems), for others it can be decadal (eg, prairie wetlands). We explore the concept of alternative stable states in unstable systems using the highly dynamic wetland ecosystems of North America's Prairie Pothole Region. The communities in these wetland ecosystems transition through multiple states in response to decadal‐long climate oscillations that cyclically influence ponded‐water depth, permanence, and chemistry. The perspective gained by considering dynamic systems in the context of stable‐state theory allows for an increased understanding of how these systems respond to changing drivers that can push them past tipping points into alternative states. Incorporation of concepts inherent to stable‐state theory has been suggested as a key scientific element upon which to base sustainable environmental management.

Alberta, Iowa, Manitoba, Minnesota, Montana, North↗

Disease-smart climate adaptation for wildlife management and conservation

Climate change is a well-documented driver and threat multiplier of infectious disease in wildlife populations. However, wildlife disease management and climate-change adaptation have largely operated in isolation. To improve conservation outcomes, we consider the role of climate adaptation in initiating or exacerbating the transmission and spread of wildlife disease and the deleterious effects thereof, as illustrated through several case studies. We offer insights into best practices for disease-smart adaptation, including a checklist of key factors for assessing disease risks early in the climate adaptation process. By assessing risk, incorporating uncertainty, planning for change, and monitoring outcomes, natural resource managers and conservation practitioners can better prepare for and respond to wildlife disease threats in a changing climate.

Frontiers in Ecology and the Environment↗

Meteorological drivers of hypolimnetic anoxia in a eutrophic, north temperate lake

Oxygen concentration is both an indicator and driver of water quality in lakes. Decreases in oxygen concentration leads to altered ecosystem function as well as harmful consequences for aquatic biota, such as fishes. The responses of oxygen dynamics in lakes to climate-related drivers, such as temperature and wind speed, are well documented for lake surface waters. However, much less is known about how the oxic environment of bottom waters, especially the timing and magnitude of anoxia in eutrophic lakes, responds to changes in climate drivers. Understanding how important ecosystem states, such as hypolimnetic anoxia, may respond to differing climate scenarios requires a model that couples physical-biological conditions and sufficiently captures the density stratification that leads to strong oxygen gradients. Here, we analyzed the effects of changes in three important meteorological drivers (air temperature, wind speed, and relative humidity) on hypolimnetic anoxia in a eutrophic, north temperate lake using the anoxic factor as an index that captures both the temporal and spatial extent of anoxia. Air temperature and relative humidity were found to have a positive correlation with anoxic factor, while wind speed had a negative correlation. Air temperature was found to have the greatest potential impact of the three drivers on the anoxic factor, followed by wind speed and then relative humidity. Across the scenarios of climate variability, variation in the simulated anoxic factor was primarily due to changes in the timing of onset and decay of stratification. Given the potential for future changes in climate, especially increases in air temperature, this study provides important insight into how these changes will alter lake water quality.

Ecological Modelling↗

Spatiotemporal dynamics of insect pollinator communities in sagebrush steppe associated with weather and vegetation

The conservation of native insect pollinators is hampered by a lack of information about environmental factors influencing pollinator communities. We investigated how insect pollinator communities, composed of bees (Hymenoptera), butterflies and moths (Lepidoptera), and flies (Diptera), are influenced by spatial and temporal aspects of the environment in sagebrush steppe shrublands. We assessed hypotheses regarding spatial autocorrelation of communities, inter-annual variability in communities, influence of elevation on timing of emergence, and influence of weather on seasonal changes in relative abundance of different pollinator taxa. We captured 27,310 insects from 6 bee families, 27 butterfly and moth families, and 3 fly families. The occurrence of insect pollinators among sampling plots was not spatially autocorrelated, indicating that insect communities may be structured by habitat variation and microclimates over relatively fine spatial scales. Pollinator familial richness, diversity, abundance, and timing of emergence were most strongly positively associated with spatiotemporal variation in minimum daily temperatures at the ground surface during the active season. Emergence timing was positively correlated with growing degree days and percent humidity, regardless of elevation. All pollinator groups varied in abundance throughout their active season, peaking in early July (bees), late July (flies), or early August (butterflies and moths). Our findings suggest that changes in nighttime temperatures, which have been steadily increasing over the last several decades as a result of climate change, may have strong effects on sagebrush steppe pollinator communities. Also, non-bee pollinators may provide particularly important pollination in this vast ecosystem during the warmest time of the year.

Oregon↗

UAV-derived models of vegetation characteristics do not transfer to extreme drought and wet conditions across a northern Arizona landscape

Context Shifts in precipitation regimes due to climate change are significantly impacting dryland ecosystems, including vegetation composition and structure. Unoccupied aerial vehicles (UAVs) are widely used to monitor vegetation, but whether models built to predict changes in these characteristics are robust under extreme precipitation regimes is unclear. Objectives We aimed to predict key vegetation characteristics under three precipitation regimes (ambient, drought, and water addition) and assess model performance across these moisture conditions. We also evaluated how models built under ambient conditions predicted vegetation characteristics under extreme precipitation regimes. Methods UAV surveys were conducted at five sites subject to long-term precipitation manipulation along an elevation gradient in northern Arizona, United States (U.S.). Twenty-one vegetation indices and point cloud data from the UAV imagery were used to develop models to predict vegetation structure and composition characteristics. Model performance and transferability were assessed via error and directional bias within each treatment (i.e., in situ) and from ambient to precipitation treatments (i.e., model transfer). Results UAV-based models accurately measured vegetation characteristics across all regimes, but maximum height showed significantly higher error under drought conditions. Models developed under ambient precipitation and applied to extreme precipitation treatments exhibited significant differences in the error and directional bias, indicating they may not be suitable under climate change. Conclusions UAV-based models are effective for monitoring vegetation characteristics but may lose accuracy under extreme precipitation regimes expected under climate change. This study emphasizes the need to improve model transferability and suggests refining landscape monitoring approaches to consider extreme changes in precipitation and associated vegetation responses.

Arizona↗

Estimating indices of range shifts in birds using dynamic models when detection is imperfect

There is intense interest in basic and applied ecology about the effect of global change on current and future species distributions. Projections based on widely used static modeling methods implicitly assume that species are in equilibrium with the environment and that detection during surveys is perfect. We used multiseason correlated detection occupancy models, which avoid these assumptions, to relate climate data to distributional shifts of Louisiana Waterthrush in the North American Breeding Bird Survey (BBS) data. We summarized these shifts with indices of range size and position and compared them to the same indices obtained using more basic modeling approaches. Detection rates during point counts in BBS surveys were low, and models that ignored imperfect detection severely underestimated the proportion of area occupied and slightly overestimated mean latitude. Static models indicated Louisiana Waterthrush distribution was most closely associated with moderate temperatures, while dynamic occupancy models indicated that initial occupancy was associated with diurnal temperature ranges and colonization of sites was associated with moderate precipitation. Overall, the proportion of area occupied and mean latitude changed little during the 1997–2013 study period. Near-term forecasts of species distribution generated by dynamic models were more similar to subsequently observed distributions than forecasts from static models. Occupancy models incorporating a finite mixture model on detection – a new extension to correlated detection occupancy models – were better supported and may reduce bias associated with detection heterogeneity. We argue that replacing phenomenological static models with more mechanistic dynamic models can improve projections of future species distributions. In turn, better projections can improve biodiversity forecasts, management decisions, and understanding of global change biology.

Global Change Biology↗

Dietary breadth of grizzly bears in the Greater Yellowstone Ecosystem

Grizzly bears ( Ursus arctos ) in the Greater Yellowstone Ecosystem (GYE) are opportunistic omnivores that eat a great diversity of plant and animal species. Changes in climate may affect regional vegetation, hydrology, insects, and fire regimes, likely influencing the abundance, range, and elevational distribution of the plants and animals consumed by GYE grizzly bears. Determining the dietary breadth of grizzly bears is important to document future changes in food resources and how those changes may affect the nutritional ecology of grizzlies. However, no synthesis exists of all foods consumed by grizzly bears in the GYE. We conducted a review of available literature and compiled a list of species consumed by grizzly bears in the GYE. We documented >266 species within 200 genera from 4 kingdoms, including 175 plant, 37 invertebrate, 34 mammal, 7 fungi, 7 bird, 4 fish, 1 amphibian, and 1 algae species as well as 1 soil type consumed by grizzly bears. The average energy values of the ungulates (6.8 kcal/g), trout (Oncorhynchus spp., 6.1 kcal/g), and small mammals (4.5 kcal/g) eaten by grizzlies were higher than those of the plants (3.0 kcal/g) and invertebrates (2.7 kcal/g) they consumed. The most frequently detected diet items were graminoids, ants (Formicidae), whitebark pine seeds (Pinus albicaulis), clover (Trifolium spp.), and dandelion (Taraxacum spp.). The most consistently used foods on a temporal basis were graminoids, ants, whitebark pine seeds, clover, elk (Cervus elaphus), thistle (Cirsium spp.), and horsetail (Equisetum spp.). Historically, garbage was a significant diet item for grizzlies until refuse dumps were closed. Use of forbs increased after garbage was no longer readily available. The list of foods we compiled will help managers of grizzly bears and their habitat document future changes in grizzly bear food habits and how bears respond to changing food resources.

Idaho, Montana, Wyoming↗

Future direction of fuels management in sagebrush rangelands

Sagebrush ecosystems in the United States have been declining since EuroAmerican settlement, largely due to agricultural and urban development, invasive species, and altered fire regimes, resulting in loss of biodiversity and wildlife habitat. To combat continued conversion to undesirable ecological states and loss of habitat to invasive species fueled by frequent fire, a variety of fuel treatments, including networks of fuel breaks, are being implemented or proposed in sagebrush ecosystems, particularly in and around the Great Basin. In this forum paper we briefly review current knowledge of common fuel treatment approaches, their intended benefits, potential risks, and limitations. We additionally discuss challenges for fuel treatment strategies in the context of changes in climate, invasive species, wildlife habitat, and human population, and we explore how advances in geospatial technologies, monitoring, and fire behavior modeling, as well as accounting for social context, can improve the efficacy of fuels management in sagebrush ecosystems. Finally, given continued potential for ecosystem transformation, we describe approaches to future fuels management by considering the applicability of the Resist-Accept-Direct (RAD) framework. The intent of the paper is to provide scientists and land managers with key information and a forward-thinking framework for fuels science and adaptive management that can respond to both expected and unexpected changes in sagebrush rangelands.

Rangeland Ecology and Management↗

Dynamic selection for forage quality and quantity in response to phenology and insects in an Arctic ungulate

Spatiotemporal variation in forage is a primary driver of ungulate behavior, yet little is known about the nutritional components they select, and how selection varies across the growing season with changes in forage quality and quantity. We addressed these uncertainties in barren-ground caribou ( Rangifer tarandus ), which experience their most important foraging opportunities during the short Arctic summer. Recent declines in Arctic caribou populations have raised concerns about the influence of climate change on summer foraging opportunities, given shifting vegetation conditions and insect harassment, and their potential effects on caribou body condition and demography. We examined Arctic caribou selection of summer forage by pairing locations from females in the Central Arctic Herd of Alaska with spatiotemporal predictions of biomass, digestible nitrogen (DN), and digestible energy (DE). We then assessed selection for these nutritional components across the growing season at landscape and patch scales, and determined whether foraging opportunities were constrained by insect harassment. During early summer, at the landscape scale, caribou selected for intermediate biomass and high DN and DE, following expectations of the forage maturation hypothesis. At the patch scale, however, caribou selected for high values of all forage components, particularly DN, suggesting that protein may be limiting. During late summer, after DN declined below the threshold for protein gain, caribou exhibited a switch at both spatial scales, selecting for higher biomass, likely enabling mass and fat deposition. Mosquito activity strongly altered caribou selection of forage and increased their movement rates, while oestrid fly activity had little influence. Our results demonstrate that early and late summer periods afford Arctic caribou distinct foraging opportunities, as they prioritize quality earlier in the summer and quantity later. Climate change may further constrain caribou access to DN as earlier, warmer Arctic summers may be associated with reduced DN and increased mosquito harassment.

Alaska↗

An aridity threshold model of fire sizes and annual area burned in extensively forested ecoregions of the western USA

Wildfire occurrence varies among regions and through time due to the long-term impacts of climate on fuel structure and short-term impacts on fuel flammability. Identifying the climatic conditions that trigger extensive fire years at regional scales can enable development of area burned models that are both spatially and temporally robust, which is crucial for understanding the impacts of past and future climate change. We identified region-specific thresholds in fire-season aridity that distinguish years with limited, moderate, and extensive area burned for 11 extensively forested ecoregions in the western United States. We developed a new area burned model using these relationships and demonstrate its application in the Southern Rocky Mountains using climate projections from five global climate models (GCMs) that bracket the range of projected changes in aridity. We used the aridity thresholds to classify each simulation year as having limited, moderate, or extensive area burned and defined fire-size distributions from historical fire records for these categories. We simulated individual fires from a regression relating fire season aridity to the annual number of fires and drew fire sizes from the corresponding fire-size distributions. We project dramatic increases in area burned after 2020 under most GCMs and after 2060 with all GCMs as the frequency of extensive fire years increases. Our adaptable model can readily incorporate new observations (e.g., extreme fire years) to directly address the non-stationarity of fire-climate relationships as climatic conditions diverge from past observations. Our aridity threshold fire model provides a simple yet spatially robust approach to project regional changes in area burned with broad applicability to ecosystem and vegetation simulation models.

Ecological Modelling↗

Low stand density moderates growth declines during hot droughts in semi-arid forests

Increasing heat and aridity in coming decades is expected to negatively impact tree growth and threaten forest sustainability in dry areas. Maintaining low stand density has the potential to mitigate the negative effects of increasingly severe droughts by minimizing competitive intensity. However, the direct impact of stand density on the growing environment (i.e. soil moisture), and the specific drought metrics that best quantify that environment, are not well explored for any forest ecosystem. We examined the relationship of varying stand density (i.e. basal area) on soil moisture and stand‐level growth in a long‐term (multi‐decadal), ponderosa pine Pinus ponderosa , forest management experiment. We accounted for the influence of stand‐level density on moisture availability by measuring and modelling soil moisture using an ecosystem water balance model. To quantify the growing environment, we developed metrics of ecological drought that integrate the influence of moisture availability in the soil with moisture demand by the atmosphere. We paired these results with stand‐level dendrochronological data, avoiding the potential bias introduced from individual tree‐based assessments, and used critical climate period analysis to identify the timing and duration of these drought metrics that most relate to forest growth. We found that stand‐level growth is highly responsive to the combination of high temperature and low soil moisture. Growth in all stands was negatively related to temperature and positively related to moisture availability, although the sensitivity of growth to those conditions varied among stand density treatments. Growth enhancement during cool years is greatest in low density stands. In addition, low density stands displayed substantially higher long‐term average growth than higher density stands and maintained higher growth even when temperatures were high. Growth in low density stands also increased more than higher density stands in response to greater long‐term moisture availability. Synthesis and applications . We quantified the influence of stand‐level density on the environmental conditions that determine tree growth and related forest growth to patterns of moisture supply and demand. Our drought metrics, and analytical approach for quantifying drought impacts on forest growth, are a novel approach for assessing forest vulnerability to drought under climate change. These results provide new perspective on the potential for density management to mitigate drought stress and maintain forest stand growth during and after drought events in water‐limited forests.

Arizona↗