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Desert shrub responses to experimental modification of precipitation seasonality and soil depth: relationship to the two-layer model and ecohydrological niche

1. Ecohydrological niches are important for understanding plant community responses to climate shifts, particularly in dry lands. According to the two-layer hypothesis, selective use of deep-soil water increases growth or persistence of woody species during warm and dry summer periods and thereby contributes to their coexistence with shallow-rooted herbs in dry ecosystems. The resource-pool hypothesis further suggests that shallow-soil water benefits growth of all plants while deep-soil water primarily enhances physiological maintenance and survival of woody species. Few studies have directly tested these by manipulating deep-soil water availability and observing the long-term outcomes. 2. We predicted that factors promoting infiltration and storage of water in deep soils, specifically greater winter precipitation and soil depth, would enhance Artemisia tridentata (big sagebrush) in cold, winter-wet/summer-dry desert. Sagebrush responses to 20 years of winter irrigation were compared to summer- or no irrigation, on plots having relatively deep or shallow soils (2 m vs. 1 m depths). 3. Winter irrigation increased sagebrush cover, and crown and canopy volumes, but not density (individuals/plot) compared to summer or no irrigation, on deep-soil plots. On shallow-soil plots, winter irrigation surprisingly decreased shrub cover and size, and summer irrigation had no effect. Furthermore, multiple regression suggested that the variations in growth were related (i) firstly to water in shallow soils (0-0.2 m) and secondly to deeper soils (> 1 m deep) and (ii) more by springtime than by midsummer soil water. Water-use efficiency increased considerably on shallow soils without irrigation and was lowest with winter irrigation. 4. Synthesis . Sagebrush was more responsive to the seasonal timing of precipitation than to total annual precipitation. Factors that enhanced deep-water storage (deeper soils plus more winter precipitation) led to increases in Artemisia tridentata that were consistent with the two-layer hypothesis, and the contribution of shallow water to growth on these plots was consistent with the resource-pool hypothesis. However, shallow-soil water also had negative effects on sagebrush, suggesting an ecohydrological trade-off not considered in these or related theories. The interaction between precipitation timing and soil depth indicates that increased winter precipitation could lead to a mosaic of increases and decreases in A. tridentata across landscapes having variable soil depth.

Idaho↗

Long-term plant responses to climate are moderated by biophysical attributes in a North American desert

Recent elevated temperatures and prolonged droughts in many already water-limited regions throughout the world, including the southwestern U.S., are likely to intensify according to future climate-model projections. This warming and drying can negatively affect perennial vegetation and lead to the degradation of ecosystem properties. To better understand these detrimental effects, we formulate a conceptual model of dryland ecosystem vulnerability to climate change that integrates hypotheses on how plant species will respond to increases in temperature and drought, including how plant responses to climate are modified by landscape, soil, and plant attributes that are integral to water availability and use. We test the model through a synthesis of fifty years of repeat measurements of perennial plant species cover in large permanent plots across the Mojave Desert, one of the most water-limited ecosystems in North America. Plant species ranged in their sensitivity to precipitation in different seasons, capacity to increase in cover with high precipitation, and resistance to decrease in cover with low precipitation. Our model successfully explains how plant responses to climate are modified by biophysical attributes in the Mojave Desert. For example, deep-rooted plants were not as vulnerable to drought on soils that allowed for deep water percolation, whereas shallow-rooted plants were better buffered from drought on soils that promoted water retention near the surface. Synthesis. Our results emphasize the importance of understanding climate-vegetation relationships in the context of biophysical attributes that influence water availability and provide an important forecast of climate-change effects, including plant mortality and land degradation in dryland regions throughout the world.

California, Nevada↗

Testing the apparent resistance of three dominant plants to chronic drought on the Colorado Plateau

Many drylands, including the south-western United States, are projected to become more water-limited as these regions become warmer and drier with climate change. Such chronic drought may push individual species or plant functional types beyond key thresholds leading to reduced growth or even mortality. Indeed, recent observational and experimental evidence from the Colorado Plateau suggests that C 3 grasses are the most vulnerable to chronic drought, while C 4 grasses and C 3 shrubs appear to have greater resistance. The effects of chronic, or press-drought are predicted to begin at the physiological level and translate up to higher hierarchical levels. To date, the drought resistance of C 4 grasses and C 3 shrubs in this region has been only evaluated at the community level and thus we lack information on whether there are sensitivities to drought at lower hierarchical levels. In this study, we tested the apparent drought resistance of three dominant species ( Pleuraphis jamesii , a C 4 rhizomatous grass; Coleogyne ramosissima , a C 3 drought-deciduous shrub; and Ephedra viridis , a C 3 evergreen shrub) to an ongoing experimental press-drought (-35% precipitation) by comparing individual-level responses (ecophysiology and growth dynamics) to community-level responses (plant cover). For all three species, we observed consistent responses across all hierarchical levels: P. jamesii was sensitive to drought across all measured variables, while the shrubs C. ramosissima and E. viridis had little to no responses to the experimental press-drought at any given level. Synthesis . Our findings suggest that the apparent drought resistance at higher hierarchical levels, such as cover, may serve as good proxies for lower-level responses. Furthermore, it appears the shrubs are avoiding drought, possibly by utilizing moisture at deeper soil layers, while the grasses are limited to shallower layers and must endure the drought conditions. Give this differential sensitivity to drought, a future with less precipitation and higher temperatures may increase the dominance of shrubs on the Colorado Plateau, as grasses succumb to chronic water stress.

Utah↗

Declines in low-elevation subalpine tree populations outpace growth in high-elevation populations with warming

Species distribution shifts in response to climate change require that recruitment increase beyond current range boundaries. For trees with long life spans, the importance of climate-sensitive seedling establishment to the pace of range shifts has not been demonstrated quantitatively. Using spatially explicit, stochastic population models combined with data from long-term forest surveys, we explored whether the climate-sensitivity of recruitment observed in climate manipulation experiments was sufficient to alter populations and elevation ranges of two widely distributed, high-elevation North American conifers. Empirically observed, warming-driven declines in recruitment led to rapid modelled population declines at the low-elevation, ‘warm edge’ of subalpine forest and slow emergence of populations beyond the high-elevation, ‘cool edge’. Because population declines in the forest occurred much faster than population emergence in the alpine, we observed range contraction for both species. For Engelmann spruce, this contraction was permanent over the modelled time horizon, even in the presence of increased moisture. For limber pine, lower sensitivity to warming may facilitate persistence at low elevations – especially in the presence of increased moisture – and rapid establishment above tree line, and, ultimately, expansion into the alpine. Synthesis . Assuming 21st century warming and no additional moisture, population dynamics in high-elevation forests led to transient range contractions for limber pine and potentially permanent range contractions for Engelmann spruce. Thus, limitations to seedling recruitment with warming can constrain the pace of subalpine tree range shifts.

Journal of Ecology↗

Growth strategies and threshold responses to water deficit modulate effects of warming on tree seedlings from forest to alpine

1.Predictions of upslope range shifts for tree species with warming are based on assumptions of moisture stress at lower elevation limits and low temperature stress at high elevation limits. However, recent studies have shown that warming can reduce tree seedling establishment across the entire gradient from subalpine forest to alpine via moisture limitation. Warming effects also vary with species, potentially resulting in community shifts in high elevation forests. 2.We examined the growth and physiology underlying effects of warming on seedling demographic patterns. We evaluated dry mass (DM), root length, allocation above- and belowground, and relative growth rate (RGR) of whole seedlings, and their ability to avoid or endure water stress via water-use efficiency and resisting turgor loss, for Pinus flexilis, Picea engelmannii and Pinus contorta seeded below, at, and above treeline in experimentally warmed, watered, and control plots in the Rocky Mountains, USA. We expected that growth and allocation responses to warming would relate to moisture status and that variation in drought tolerance traits would explain species differences in survival rates. 3.Across treatments and elevations, seedlings of all species had weak turgor-loss resistance, and growth was marginal with negative RGR in the first growth phase (-0.01 to -0.04 g/g/d). Growth was correlated with soil moisture, particularly in the relatively small-seeded P. contorta and P. engelmannii. P. flexilis, known to have the highest survivorship, attained the greatest DM and longest root but was also the slowest growing and most water-use-efficient. This was likely due to its greater reliance on seed reserves. Seedlings developed 15% less total DM, 25% less root DM, and 11% shorter roots in heated compared to unheated plots. Higher temperatures slightly increased DM, root length and RGR where soils were wettest, but more strongly decreased these variables under drier conditions. 4.Synthesis: The surprising heat-inhibition of tree seedling establishment at the cold edge of forests appears to have a physiological basis: newly germinated seedlings have poor moisture stress tolerance, which appears related to marginal initial growth and heavy reliance on seed reserves. Variation in these attributes among tree species at treeline helps explain their different climate responses.

Journal of Ecology↗

Ecosystem vs. community recovery 25 years after grass invasions and fire in a subtropical woodland

Despite a large body of research documenting invasive plant impacts, few studies have followed individual invaded sites over decades to observe how they change, and none have contrasted how compositional impacts from invasion compare to ecosystem-process impacts over a multi-decadal time-scale. Using direct measurements of plant density and composition and of ecosystems processes, we evaluate how ecosystem structure, above-ground net primary production (ANPP), and above-ground and soil nutrient pools compare over 25 years since fire and C4 grass invasions disrupted seasonally dry Hawaiian woodlands. We compare structure and function between primary woodland that has never burned and is largely native species-dominated, with sites that had been the same woodland type but burned in alien-grass-fuelled fires in the 1970s and 1980s. The sites have not experienced fires since 1987. We report here that woody plant composition and structure continue to be dramatically changed by the initial invasions and fires that occurred 25 years ago and invaders continue to dominate in burned sites. This is reflected in continued low plant carbon pools in burned compared to unburned sites. Yet ANPP and N storage, which were dramatically lower in the initial decade after invasive-grass fuelled fires, have increased and are now indistinguishable from values measured in intact woodlands. Soil carbon pools were resilient to both invasion and fire initially and over time. Above-ground net primary production has recovered because of invasion of burned sites by a non-native N-fixing tree rather than because of recovery of native species. This invasive N-fixing tree is unlikely to return C storage of the invaded sites to those of unburned woodland because of its tissue and growth characteristics and its interactions with invasive grasses. It does not facilitate native species but rather promotes a persistent invasive grass/N-fixer savanna. Synthesis. We conclude that fire, an unusual disturbance in this system, has perpetuated the dominance of these sites by invasive species and that despite the dramatic recovery of above-ground net primary production and N pools, the ecosystem continues to be in a distinctly different state than the pre-fire, pre-Melinis community. Thus, despite the absence of further disturbance (fire), there is no evidence that succession towards the original ecosystem is occurring. The fact that N pools and above-ground net primary production recover because of a new invader (Morella faya), highlights the unpredictability of ecosystem trajectories in the face of altered regional species pools.

Hawaii↗

Alternative pathways to landscape transformation: Invasive grasses, burn severity and fire frequency in arid ecosystems

Arid ecosystems are often vulnerable to transformation to invasive-dominated states following fire, but data on persistence of these states are sparse. The grass/fire cycle is a feedback process between invasive annual grasses and fire frequency that often leads to the formation of alternative vegetation states dominated by the invasive grasses. However, other components of fire regimes, such as burn severity, also have the potential to produce long-term vegetation transformations. Our goal was to evaluate the influence of both fire frequency and burn severity on the transformation of woody-dominated communities to communities dominated by invasive grasses in major elevation zones of the Mojave Desert of western North America. We used a chronosequence design to collect data on herbaceous and woody cover at 229 unburned reference plots and 578 plots that burned between 1972 and 2010. We stratified the plots by elevation zone (low, mid, high), fire frequency (1–3 times) and years post-fire (YPF; 1–5, 6–10, 11–20 and 21–40 YPF). Burn severity for each plot was estimated by the difference normalized burn ratio. We identified two broad post-fire successional pathways. One was an outcome of fire frequency, resulting in a strong potential transformation via the grass/fire cycle. The second pathway was driven by burn severity, the critical aspect being that long-term transformation of a community could occur from just one fire in areas that burned at high or sometimes moderate severity. Dominance by invasive grasses was most likely to occur in low-and high-elevation communities; cover of native herbaceous species was often greater than that of invasive grasses in the mid-elevation zone. Synthesis . Invasive grasses can dominate a site that burned only one time in many decades at high severity, or a site that burned at low severity but multiple times in the same time period. However, high burn severity may predispose areas to more frequent fire because they have relatively high cover of invasive annual grass, suggesting burn severity and fire frequency have both independent and synergistic effects. Resilience in vegetation structure following fire in many arid communities may be limited to a narrow window of low burn severity in areas that have not burned in many decades.

Mojave Desert↗

Effect of permafrost thaw on plant and soil fungal community in the boreal forest: Does fungal community change mediate plant productivity response?

Permafrost thaw is leading to rapid shifts in boreal ecosystem function. Permafrost thaw affects soil carbon turnover through changes in soil hydrology, however, the biotic mechanisms regulating plant community response remain elusive. Here, we measured the response of fungal community composition and soil nutrient content in an intact permafrost plateau forest soil and an adjacent thermokarst bog using barcoded amplicon targeting ITS2 and 28S rRNA genes. Next, we used the soils from the permafrost plateau and the thermokarst bog as soil inoculum in a greenhouse experiment to measure whether shifts in fungal community and soil water level regulate plant productivity. Overall, we found that fungal community composition differed significantly between the thawed and intact permafrost sites, but soil nutrient content did not. Relative abundance of mycorrhizal fungal taxa decreased while relative abundance of putative fungal pathogens increased with permafrost thaw. In the greenhouse, we found that ecto- and arbuscular associated host plants had higher productivity in permafrost-intact soils relative to thawed soils. However, productivity of non-mycorrhizal tussock grass was more affected by soil water levels than soil communities. Our results suggest that fungal communities are crucial in mediating plant response to permafrost thaws inducing hydrology changes.

Alaska↗

Which trees die during drought? The key role of insect host-tree selection

1. During drought, the tree subpopulations (such as size or vigor classes) that suffer disproportionate mortality can be conceptually arrayed along a continuum defined by the actions of biotic agents, particularly insects. At one extreme, stress dominates: insects are absent or simply kill the most physiologically stressed trees. At the opposite extreme, host selection dominates: outbreaking insects kill trees independently of their stress, instead selecting trees based on size or other traits. Intermediate responses are also possible. Yet for mixed-species forests, we lack a broad understanding of the relative importance of insects in determining exactly which subpopulations of trees suffer disproportionate mortality during drought, and whether these subpopulations differ among co-occurring tree species. 2. During an extreme drought, we documented the roles of native bark beetles in the mortality of five tree species in California’s Sierra Nevada. We analyzed patterns and agents of tree mortality in 12 permanent plots, and patterns of mortality in 89 temporary plots. 3. Most tree mortality was associated with bark beetles. But the growth rates (an indicator of chronic stress) and sizes of trees that suffered greatest bark-beetle-related mortality differed sharply among tree taxa, variously conforming with domination by stress (Abies concolor), domination by host selection (Pinus lambertiana and P. ponderosa), or a mix of the two (Calocedrus decurrens). Quercus kelloggii mortality remained relatively low. Thus, even during extreme drought substantial proportions of stressed trees survived because they were of sizes that mostly avoided fatal insect attack. Conversely, substantial proportions of comparatively unstressed trees died because they were of sizes that were selectively killed by outbreaking insects. 4. Synthesis. Native bark beetles were primarily responsible for determining which subpopulations of trees suffered greatest mortality during drought. However, idiosyncratic host-tree selection by the different bark beetle taxa meant that the tree subpopulations suffering greatest mortality differed strikingly among tree taxa – for example, high mortality of small trees of one species, but of large trees of another. If idiosyncratic host-tree selection by biotic mortality agents proves to be a generally common phenomenon, it could help explain weak broad-scale correlations between tree traits and tree mortality during drought.

Journal of Ecology↗

Phenological mismatch between season advancement and migration timing alters Arctic plant traits

1. Climate change is creating phenological mismatches between herbivores and their plant resources throughout the Arctic. While advancing growing seasons and changing arrival times of migratory herbivores has been shown to have consequences for herbivores and forage quality, developing mismatches are also likely to influence other traits of plants, such as above- and belowground biomass and the type of reproduction, that are often not investigated. 2. In coastal western Alaska, we conducted a three-year factorial experiment that simulated scenarios of phenological mismatch by manipulating the start of the growing season (ca. 3-weeks early and ambient) and grazing times (3-weeks early, typical, 3-weeks late, or no-grazing) of Pacific black brant (Branta bernicla nigricans), to examine how the timing of these events influence a primary goose forage species, Carex subspathacea. 3. After three years, an advanced growing season compared to a typical growing season increased stem heights, standing dead biomass, and the number of inflorescences. Early season grazing compared to typical season grazing reduced above- and belowground biomass, stem height, and the number of tillers; while late season grazing increased the number of inflorescences and standing dead biomass by year 3. Therefore, an advanced growing season and late grazing had comparable directional effects on most plant traits, but a 3-week delay in grazing had an impact on vegetation traits 3 to 5 times greater than a similar shift in advancement of spring conditions. In addition, changes in response to treatments for some variables, such as the number of inflorescences, were not measurable until the second year of the experiment, while other variables, such as root productivity and number of tillers, changed the direction of their responses to treatments over time. 4. Synthesis: Factors influencing the timing of migration have a larger influence on an important forage species than earlier springs in the breeding and rearing habitats of Pacific black brant. The phenological mismatch prediction for this site of earlier springs and later goose arrival will likely increase above- and belowground biomass and sexual reproduction of C. subspathacea. Implications for mismatch may be difficult to predict because some variables required successive years of mismatch to respond.

Journal of Ecology↗

Frequent burning causes large losses of carbon from deep soil layers in a temperate savanna

Fire activity is changing dramatically across the globe, with uncertain effects on ecosystem processes, especially below‐ground. Fire‐driven losses of soil carbon (C) are often assumed to occur primarily in the upper soil layers because the repeated combustion of above‐ground biomass limits organic matter inputs into surface soil. However, C losses from deeper soil may occur if frequent burning reduces root biomass inputs of C into deep soil layers or stimulates losses of C via leaching and priming. To assess the effects of fire on soil C, we sampled 12 plots in a 51‐year‐long fire frequency manipulation experiment in a temperate oak savanna, where variation in prescribed burning frequency has created a gradient in vegetation structure from closed‐canopy forest in unburned plots to open‐canopy savanna in frequently burned plots. Soil C stocks were nonlinearly related to fire frequency, with soil C peaking in savanna plots burned at an intermediate fire frequency and declining in the most frequently burned plots. Losses from deep soil pools were significant, with the absolute difference between intermediately burned plots versus most frequently burned plots more than doubling when the full 1 m sample was considered rather than the top 0–20 cm alone (losses of 98.5 Mg C/ha [−76%] and 42.3 Mg C/ha [−68%] in the full 1 m and 0–20 cm layers respectively). Compared to unburned forested plots, the most frequently burned plots had 65.8 Mg C/ha (−58%) less C in the full 1 m sample. Root biomass below the top 20 cm also declined by 39% with more frequent burning. Concurrent fire‐driven losses of nitrogen and gains in calcium and phosphorus suggest that burning may increase nitrogen limitation and play a key role in the calcium and phosphorus cycles in temperate savannas. Synthesis . Our results illustrate that fire‐driven losses in soil C and root biomass in deep soil layers may be critical factors regulating the net effect of shifting fire regimes on ecosystem C in forest‐savanna transitions. Projected changes in soil C with shifting fire frequencies in savannas may be 50% too low if they only consider changes in the topsoil.

Minnesota↗

More than one way to kill a spruce forest: The role of fire and climate in the late-glacial termination of spruce woodlands across the southern Great Lakes

In the southern Great Lakes Region, North America, between 19,000 and 8,000 years ago, temperatures rose by 2.5–6.5°C and spruce Picea forests/woodlands were replaced by mixed-deciduous or pine Pinus forests. The demise of Picea forests/woodlands during the last deglaciation offers a model system for studying how changing climate and disturbance regimes interact to trigger declines of dominant species and vegetation-type conversions. The role of rising temperatures in driving the regional demise of Picea forests/woodlands is widely accepted, but the role of fire is poorly understood. We studied the effect of changing fire activity on Picea declines and rates of vegetation composition change using fossil pollen and macroscopic charcoal from five high-resolution lake sediment records. The decline of Picea forests/woodlands followed two distinct patterns. At two sites (Stotzel-Leis and Silver Lake), fire activity reached maximum levels during the declines and both charcoal accumulation rates and fire frequency were significantly and positively associated with vegetation composition change rates. At these sites, Picea declined to low levels by 14 kyr BP and was largely replaced by deciduous hardwood taxa like ash Fraxinus , hop-hornbeam/hornbeam Ostrya/Carpinus and elm Ulmus . However, this ecosystem transition was reversible, as Picea re-established at lower abundances during the Younger Dryas. At the other three sites, there was no statistical relationship between charcoal accumulation and vegetation composition change rates, though fire frequency was a significant predictor of rates of vegetation change at Appleman Lake and Triangle Lake Bog. At these sites, Picea declined gradually over several thousand years, was replaced by deciduous hardwoods and high levels of Pinus and did not re-establish during the Younger Dryas. Synthesis . Fire does not appear to have been necessary for the climate-driven loss of Picea woodlands during the last deglaciation, but increased fire frequency accelerated the decline of Picea in some areas by clearing the way for thermophilous deciduous hardwood taxa. Hence, warming and intensified fire regimes likely interacted in the past to cause abrupt losses of coniferous forests and could again in the coming decades.

Indiana, Michigan, Ohio↗

Stress gradients interact with disturbance to reveal alternative states in salt marsh: Multivariate resilience at the landscape scale

Stress gradients influence many ecosystem processes and properties, including ecosystem recovery from and resistance to disturbance. While recent analytical approaches have advanced multivariate metrics of ecosystem resilience that allow quantification of conceptual resilience models and identification of thresholds of state change, these approaches are not often translated to landscape scales. Using natural and restored salt marshes in Louisiana, USA, we quantified plant community recovery and resistance metrics along flooding stress gradients. n ‐dimensional hypervolumes of plant community biomass and structure were simulated using field data collected from disturbance‐recovery experiments. The relationships between multivariate resilience metrics and flooding stress gradients were then mapped at community‐ and landscape‐relevant scales by scaling with airborne‐derived data across the region. Greater pre‐disturbance abiotic stress decreased live belowground, but not aboveground, biomass, and ultimately led to lower post‐disturbance total recovery, recovery rates, and resistance of plant communities. Vegetated plots flooded >52% of the time transitioned to an alternative, unvegetated state after disturbance. Mapping revealed differences in spatial patterns of resilience‐ highlighting low, interior marsh edges as especially vulnerable to the combination of chronic flooding stress and acute disturbance. At the landscape scale, approximately half of the area (48%) is vulnerable to state change after pulse disturbances. Synthesis . Ultimately, we quantify the ball‐and‐cup conceptual model for a salt marsh ecosystem and its alternative state, mudflat. We find that increasing abiotic stress due to climate change diminishes ecosystem resilience, but the interaction with common episodic disturbances is necessary to reveal transitions to alternative states and quantify state change thresholds. Quantifying and mapping resilience and where alternative states may exist in this fashion improves ecologists’ ability to investigate the mechanisms of stress gradient control on emergent ecosystem properties, while providing spatially explicit resources for managing ecosystems according to their projected resilience.

Louisiana↗

Increased burning in a warming climate reduces carbon uptake in the Greater Yellowstone Ecosystem despite productivity gains

1. The effects of changing climate and disturbance on mountain forest carbon stocks vary with tree species distributions and over elevational gradients. Warming can increase carbon uptake by stimulating productivity at high elevations but also enhance carbon release by increasing respiration and the frequency, intensity, and size of wildfires. 2. To understand the consequences of climate change for temperate mountain forests, we simulated interactions among climate, wildfire, tree species, and their combined effects on regional carbon stocks in forests of the Greater Yellowstone Ecosystem, USA with the LANDIS‐II landscape change model. Simulations used historical climate and future potential climate represented by downscaled projections from five general circulation models (GCMs) that bracket the range of variability under the representative concentration pathway (RCP) 8.5 emissions scenario. 3. Total ecosystem carbon increased by 67% through 2100 in simulations with historical climate, and by 38 – 69% with GCM climate. Differences in carbon uptake among GCMs resulted primarily from variation in area burned, not productivity. Warming increased productivity by extending the growing season, especially near upper treeline, but did not offset biomass losses to fire. By 2100, simulated area burned increased by 27 – 215% under GCM climate, with the largest increases after 2050. With warming >3 °C in mean annual temperature, the increased frequency of large fires reduced live carbon stocks by 4 – 36% relative to the control, historical climate scenario. However, relative losses in total carbon were delayed under GCMs with large increases in summer precipitation and buffered by carbon retained in soils and the wood of fire‐killed trees. Increasing fire size limited seed dispersal, and reductions in soil moisture limited seedling establishment; both effects will likely constrain long‐term forest regeneration and carbon uptake. 4. Synthesis. Forests in the GYE can maintain a carbon sink through the mid‐century in a warming climate but continued warming may cause the loss of forest area, live aboveground biomass, and ultimately, ecosystem carbon. Future changes in carbon stocks in similar forests throughout western North America will depend on regional thresholds for extensive wildfire and forest regeneration and therefore, changes may occur earlier in drier regions.

Idaho, Montana, Wyoming↗

Weather and distance to fire refugia limit landscape‐level occurrence of fungal disease in an exotic annual grass

The enemy release hypothesis proposes that invasion by exotic plant species is driven by their release from natural enemies (i.e. herbivores and pathogens) in their introduced ranges. However, in many cases, natural enemies, which may be introduced or managed to regulate invasive species, may fail to impact target host populations. Landscape heterogeneity, which can affect both the population dynamics of the pathogen and the susceptibility and the density of hosts, may contribute to why pathogens fail to control hosts despite established negative disease impacts. We explored patterns of post‐fire infection of the fungal head‐smut pathogen Ustilago bullata on the invasive annual cheatgrass Bromus tectorum , which has caused the notorious grass‐fire cycle and ecosystem degradation across Western North America. We asked whether infection level was a driver of host density or vice‐versa, and how weather affected infection and how spatial patterns of infection varied with time since fire, using a combination of structural equation modelling (SEM), proportional odds modelling and entropy‐based local indicator of spatial association (ELSA) on data from >700 plots spanning >100,000 ha remeasured annually for 4 years. Observed infection levels increased with greater prior‐year cheatgrass cover, and disease severity did not suppress cheatgrass populations. Warm, humid fall/winters and proximity to fire refugia (unburned patches) were associated with more infections. Infection clustering was most evident 2–3 years following fire with warm‐wet fall–winter conditions and decreased after two drier, colder winters. Synthesis . Severity of fungal disease did not result in measurable reductions of populations of a non‐native, invasive host species, cheatgrass, which suggests that natural enemies may not strongly regulate cheatgrass in its introduced range. Landscape heterogeneity associated with disturbance and weather limited population‐level infection of hosts by the fungal pathogen. Disturbance (specifically wildfire) and variable weather are key components of this and similar invasion systems, and likely need to be considered when evaluating disease dynamics and potential for natural enemies to influence invasion potential.

Journal of Ecology↗

Species mixture effects and climate influence growth, recruitment and mortality in Interior West USA Populus tremuloides-conifer communities

Tree-species mixture effects (e.g. complementarity and facilitation) have been found to increase individual-tree productivity, lessen mortality and improve recruitment in forests worldwide. By promoting more efficient and complete resource use, mixture effects may also lessen individual-tree-level water stress, thus improving drought resistance. We investigated the influence of mixture effects on tree productivity, mortality and recruitment across broad compositional and moisture gradients in high-elevation Interior West US mixed-conifer communities, where Populus tremuloides (trembling aspen) is the major contributor to functional diversity. Our goal was to provide a more complete scientific foundation for managing these drought-prone, fire-excluded systems under an uncertain climate. We used landscape-scale national forest inventory data to examine mixture effects on P. tremuloides and the major associated conifer species, Pseudotsuga menziesii , Pinus contorta , Abies lasiocarpa and Picea engelmannii . Using generalized linear mixed modelling, we isolated the influences of P. tremuloides relative density and climate on tree-level (stems ≥ 12.7 cm DBH) growth, mortality and stand-level recruitment (presence/absence of new trees). Cold-season precipitation (PPT) and warm-season vapour pressure deficit (VPD) served to represent soil moisture supply and demand, respectively. Populus tremuloides growth declined as interspecific density increased. In contrast, Pinus contorta and A. lasiocarpa growth increased with P. tremuloides density. For all species except A. lasiocarpa and P. menziesii, growth increased under higher PPT and VPD. Populus tremuloides mortality increased under high VPD but not with interspecific relative density. We found limited evidence that A. lasiocarpa mortality decreased as P. tremuloides density increased. Populus tremuloides recruitment declined steeply above 25% interspecific relative density. We found a decline in conifer recruitment odds as P. tremuloides density increased, ranging from strong in P. contorta to insubstantial in P. engelmannii . Synthesis . Our findings have implications for sustaining mixed-conifer communities impacted by climate change and historical fire exclusion. Mixtures of P. tremuloides and conifers may improve conifer growth while adversely impacting P. tremuloides growth relative to pure stands. Higher conifer productivity combined with lower P. tremuloides recruitment at high conifer relative density may accelerate succession.

Journal of Ecology↗

Local climate adaptations in two ubiquitous Mojave Desert shrub species, Ambrosia dumosa and Larrea tridentata

Widely distributed species are often locally adapted to climate gradients across their ranges. But little is known about the patterns of intraspecific adaptation in desert shrubs. We examined the questions of local adaptation in multiple populations of two common shrub species of the winter-wet Mojave Desert in North America in a multiple common garden experiment. Plants were raised in the greenhouse and transplanted at the age of 1 year. Ambrosia dumosa is a drought-deciduous low shrub and Larrea tridentata is an exceptionally long-lived evergreen. Over 4 years, we monitored growth, survivorship, leaf and reproductive cover and once measured leaf N content, δ 13 C and SLA. We hypothesized that populations of both species would be differentiated along a growth–survivorship trade-off according to homesite aridity. Both species exhibited previously undocumented population differences along gradients of winter precipitation and temperature. In general, populations from more winter-mesic regions had faster growth in more mesic gardens and lower survivorship in the most arid garden. Homesites with more variable summer precipitation had greater growth for A. dumosa populations, but lower growth for L. tridentata . Among L. tridentata populations, leaf cover correlated positively with growth and negatively with survival time. For A. dumosa populations, growth and survival could not be attributed to specific traits across gardens. However, larger transplants had generally lower growth rates and higher survival rates across gardens, except in the driest garden, where the population averages of intrinsic water use efficiency (iWUE) and stem growth rate were positively correlated. Synthesis . Two dominant species of the Mojave Desert adapted locally to variation in winter and summer precipitation and temperature. They did so in different ways, suggesting that L. tridentata mitigated the risk of hydraulic failure, while A. dumosa optimized carbon assimilation for growth.

California, Nevada↗