Dryland soil recovery after disturbance across soil and climate gradients of the Colorado Plateau.

Drylands impacted by energy development often require costly reclamation activities to reconstruct damaged soils and vegetation, yet little is known about the effectiveness of reclamation practices in promoting recovery of soil quality due to a lack of long-term and cross-site studies. Here, we examined paired on-pad and adjacent undisturbed off-pad soil properties over a 22-year chronosequence of 91 reclaimed oil or gas well pads across soil and climate gradients of the Colorado Plateau in the southwestern United States. Our goals were to estimate the time required for soil properties to reach undisturbed conditions, examine the multivariate nature of soil quality following reclamation, and identify environmental factors that affect reclamation outcomes. Soil samples, collected in 2020 and 2021, were analyzed for biogeochemical pools (total nitrogen, and total organic and inorganic carbon), chemical characteristics (salinity, sodicity, pH), and texture. Predicted time to recovery across all sites was 29 years for biogeochemical soil properties, 31 years for soil chemical properties, and 6 years for soil texture. Ordination of soil properties revealed differences between on- and off-pad soils, while site aridity explained variability in on-pad recovery. The predicted time to total soil recovery (distance between on- and off-pad in ordination space) was 96 years, which was longer than any individual soil property. No site reached total recovery, indicating that individual soil properties alone may not fully indicate recovery in soil quality as soil recovery does not equal the sum of its parts. Site aridity was the largest predictor of reclamation outcomes, but the effects differed depending on soil type. Taken together, results suggest the recovery of soil quality - which reflects soil fertility, carbon sequestration potential, and other ecosystem functions - was influenced primarily by site setting, with soil type and aridity major mediators of on-pad carbon, salinity, and total soil recovery following reclamation.

A conserved interdomain microbial network underpins cadaver decomposition despite environmental variables.

Microbial breakdown of organic matter is one of the most important processes on Earth, yet the controls of decomposition are poorly understood. Here we track 36 terrestrial human cadavers in three locations and show that a phylogenetically distinct, interdomain microbial network assembles during decomposition despite selection effects of location, climate and season. We generated a metagenome-assembled genome library from cadaver-associated soils and integrated it with metabolomics data to identify links between taxonomy and function. This universal network of microbial decomposers is characterized by cross-feeding to metabolize labile decomposition products. The key bacterial and fungal decomposers are rare across non-decomposition environments and appear unique to the breakdown of terrestrial decaying flesh, including humans, swine, mice and cattle, with insects as likely important vectors for dispersal. The observed lockstep of microbial interactions further underlies a robust microbial forensic tool with the potential to aid predictions of the time since death.

Trajectories and tipping points of piñon-juniper woodlands after fire and thinning.

Piñon-juniper (PJ) woodlands are a dominant community type across the Intermountain West, comprising over a million acres and experiencing critical effects from increasing wildfire. Large PJ mortality and regeneration failure after catastrophic wildfire have elevated concerns about the long-term viability of PJ woodlands. Thinning is increasingly used to safeguard forests from fire and in an attempt to increase climate resilience. We have only a limited understanding of how fire and thinning will affect the structure and function of PJ ecosystems. Here, we examined vegetation structure, microclimate conditions, and PJ regeneration dynamics following ~20 years post-fire and thinning treatments. We found that burned areas had undergone a state shift that did not show signs of returning to their previous state. This shift was characterized by (1) distinct plant community composition dominated by grasses; (2) a lack of PJ recruitment; (3) a decrease in the sizes of interspaces in between plants; (4) lower abundance of late successional biological soil crusts; (5) lower mean and minimum daily soil moisture values; (6) lower minimum daily vapor pressure deficit; and (7) higher photosynthetically active radiation. Thinning created distinct plant communities and served as an intermediate between intact and burned communities. More intensive thinning decreased PJ recruitment and late successional biocrust cover. Our results indicate that fire has the potential to create drier and more stressful microsite conditions, and that, in the absence of active management following fire, there may be shifts to persistent ecological states dominated by grasses. Additionally, more intensive thinning had a larger impact on community structure and recruitment than less intensive thinning, suggesting that careful consideration of goals could help avoid unintended consequences. While our results indicate the vulnerability of PJ ecosystems to fire, they also highlight management actions that could be adapted to create conditions that promote PJ re-establishment.

Cultivating Resilience in Dryland Soils: An Assisted Migration Approach to Biological Soil Crust Restoration.

Land use practices and climate change have driven substantial soil degradation across global drylands, impacting ecosystem functions and human livelihoods. Biological soil crusts, a common feature of dryland ecosystems, are under extensive exploration for their potential to restore the stability and fertility of degraded soils through the development of inoculants. However, stressful abiotic conditions often result in the failure of inoculation-based restoration in the field and may hinder the long-term success of biocrust restoration efforts. Taking an assisted migration approach, we cultivated biocrust inocula sourced from multiple hot-adapted sites (Mojave and Sonoran Deserts) in an outdoor facility at a cool desert site (Colorado Plateau). In addition to cultivating inoculum from each site, we created an inoculum mixture of biocrust from the Mojave Desert, Sonoran Desert, and Colorado Plateau. We then applied two habitat amelioration treatments to the cultivation site (growth substrate and shading) to enhance soil stability and water availability and reduce UV stress. Using marker gene sequencing, we found that the cultivated mixed inoculum comprised both local- and hot-adapted cyanobacteria at the end of cultivation but had similar cyanobacterial richness as each unmixed inoculum. All cultivated inocula had more cyanobacterial 16S rRNA gene copies and higher cyanobacterial richness when cultivated with a growth substrate and shade. Our work shows that it is possible to field cultivate biocrust inocula sourced from different deserts, but that community composition shifts toward that of the cultivation site unless habitat amelioration is employed. Future assessments of the function of a mixed inoculum in restoration and its resilience in the face of abiotic stressors are needed to determine the relative benefit of assisted migration compared to the challenges and risks of this approach.

Thermography captures the differential sensitivity of dryland functional types to changes in rainfall event timing and magnitude.

Drylands of the southwestern United States are rapidly warming, and rainfall is becoming less frequent and more intense, with major yet poorly understood implications for ecosystem structure and function. Thermography-based estimates of plant temperature can be integrated with air temperature to infer changes in plant physiology and response to climate change. However, very few studies have evaluated plant temperature dynamics at high spatiotemporal resolution in rainfall pulse-driven dryland ecosystems. We address this gap by incorporating high-frequency thermal imaging into a field-based precipitation manipulation experiment in a semi-arid grassland to investigate the impacts of rainfall temporal repackaging. All other factors held constant, we found that fewer/larger precipitation events led to cooler plant temperatures (1.4°C) compared to that of many/smaller precipitation events. Perennials, in particular, were 2.5°C cooler than annuals under the fewest/largest treatment. We show these patterns were driven by: increased and consistent soil moisture availability in the deeper soil layers in the fewest/largest treatment; and deeper roots of perennials providing access to deeper plant available water. Our findings highlight the potential for high spatiotemporal resolution thermography to quantify the differential sensitivity of plant functional groups to soil water availability. Detecting these sensitivities is vital to understanding the ecohydrological implications of hydroclimate change.

Assessing community assembly controls over community-scale nutrient resorption responses to nitrogen deposition.

Nutrient resorption is a fundamental physiological process in plants, with important ecological controls over numerous ecosystem functions. However, the role of community assembly in driving responses of nutrient resorption to perturbation remains largely unknown. Following the Price equation framework and the Community Assembly and Ecosystem Function framework, we quantified the contribution of species loss, species gain, and shared species to the reduction of community-level nutrient resorption efficiency in response to multi-level nitrogen (N) addition in a temperate steppe, after continuous N addition for seven years. Reductions of both N and phosphorus (P) resorption efficiency (NRE and PRE, respectively) were positively correlated with N addition levels. The dissimilarities in species composition between N-enriched and control communities increased with N addition levels, and N-enriched plots showed substantial species losses and gains. Interestingly, the reduction of community-scale NRE and PRE mostly resulted from N-induced decreases in resorption efficiency for the shared species in the control and N-enriched communities. There were negative correlations between the contributions of species richness effect and species identity effect and between the number and identity of species gained for the changes in both NRE and PRE following N enrichment. By simultaneously considering N-induced changes in species composition and in species-level resorption, our work presents a more complete picture of how different community assembly processes contribute to N-induced changes in community-level resorption.

Soil surface treatments and precipitation timing determine seedling development across southwestern US restoration sites.

Restoration in dryland ecosystems often has poor success due to low and variable water availability, degraded soil conditions, and slow plant community recovery rates. Restoration treatments can mitigate these constraints but, because treatments and subsequent monitoring are typically limited in space and time, our understanding of their applicability across broader environmental gradients remains limited. To address this limitation, we implemented and monitored a standardized set of seeding and soil surface treatments (pits, mulch, and ConMod artificial nurse plants) designed to enhance soil moisture and seedling establishment across RestoreNet, a growing network of 21 diverse dryland restoration sites in the southwestern USA over 3 years. Generally, we found that the timing of precipitation relative to seeding and the use of soil surface treatments were more important in determining seeded species emergence, survival, and growth than site-specific characteristics. Using soil surface treatments in tandem with seeding promoted up to 3× greater seedling emergence densities compared with seeding alone. The positive effect of soil surface treatments became more prominent with increased cumulative precipitation since seeding. The seed mix type with species currently found within or near a site and adapted to the historical climate promoted greater seedling emergence densities compared with the seed mix type with species from warmer, drier conditions expected to perform well under climate change. Seed mix and soil surface treatments had a diminishing effect as plants developed beyond the first season of establishment. However, we found strong effects of the initial period seeded and of the precipitation leading up to each monitoring date on seedling survival over time, especially for annual and perennial forbs. The presence of exotic species exerted a negative influence on seedling survival and growth, but not initial emergence. Our findings suggest that seeded species recruitment across drylands can generally be promoted, regardless of location, by (1) incorporation of soil surface treatments, (2) employment of near-term seasonal climate forecasts, (3) suppression of exotic species, and (4) seeding at multiple times. Taken together, these results point to a multifaceted approach to ameliorate harsh environmental conditions for improved seeding success in drylands, both now and under expected aridification.

Droughting a megadrought: Ecological consequences of a decade of experimental drought atop aridification on the Colorado Plateau.

Global dryland vegetation communities will likely change as ongoing drought conditions shift regional climates towards a more arid future. Additional aridification of drylands can impact plant and ground cover, biogeochemical cycles, and plant-soil feedbacks, yet how and when these crucial ecosystem components will respond to drought intensification requires further investigation. Using a long-term precipitation reduction experiment (35% reduction) conducted across the Colorado Plateau and spanning 10 years into a 20+ year regional megadrought, we explored how vegetation cover, soil conditions, and growing season nitrogen (N) availability are impacted by drying climate conditions. We observed large declines for all dominant plant functional types (C3 and C4 grasses and C3 and C4 shrubs) across measurement period, both in the drought treatment and control plots, likely due to ongoing regional megadrought conditions. In experimental drought plots, we observed less plant cover, less biological soil crust cover, warmer and drier soil conditions, and more soil resin-extractable N compared to the control plots. Observed increases in soil N availability were best explained by a negative correlation with plant cover regardless of treatment, suggesting that declines in vegetation N uptake may be driving increases in available soil N. However, in ecosystems experiencing long-term aridification, increased N availability may ultimately result in N losses if soil moisture is consistently too dry to support plant and microbial N immobilization and ecosystem recovery. These results show dramatic, worrisome declines in plant cover with long-term drought. Additionally, this study highlights that more plant cover losses are possible with further drought intensification and underscore that, in addition to large drought effects on aboveground communities, drying trends drive significant changes to critical soil resources such as N availability, all of which could have long-term ecosystem impacts for drylands.

Patterns and controls of foliar nutrient stoichiometry and flexibility across United States forests.

Plant element stoichiometry and stoichiometric flexibility strongly regulate ecosystem responses to global change. Here, we tested three potential mechanistic drivers (climate, soil nutrients, and plant taxonomy) of both using paired foliar and soil nutrient data from terrestrial forested National Ecological Observatory Network sites across the USA. We found that broad patterns of foliar nitrogen (N) and foliar phosphorus (P) are explained by different mechanisms. Plant taxonomy was an important control over all foliar nutrient stoichiometries and concentrations, especially foliar N, which was dominantly related to taxonomy and did not vary across climate or soil gradients. Despite a lack of site-level correlations between N and environment variables, foliar N exhibited intraspecific flexibility, with numerous species-specific correlations between foliar N and various environmental factors, demonstrating the variable spatial and temporal scales on which foliar chemistry and stoichiometric flexibility can manifest. In addition to plant taxonomy, foliar P and N:P ratios were also linked to soil nutrient status (extractable P) and climate, especially actual evapotranspiration rates. Our findings highlight the myriad factors that influence foliar chemistry and show that broad patterns cannot be explained by a single consistent mechanism. Furthermore, differing controls over foliar N versus P suggests that each may be sensitive to global change drivers on distinct spatial and temporal scales, potentially resulting in altered ecosystem N:P ratios that have implications for processes ranging from productivity to carbon sequestration.

The consequences of climate change for dryland biogeochemistry.

Drylands, which cover > 40% of Earth's terrestrial surface, are dominant drivers of global biogeochemical cycling and home to more than one third of the global human population. Climate projections predict warming, drought frequency and severity, and evaporative demand will increase in drylands at faster rates than global means. As a consequence of extreme temperatures and high biological dependency on limited water availability, drylands are predicted to be exceptionally sensitive to climate change and, indeed, significant climate impacts are already being observed. However, our understanding and ability to forecast climate change effects on dryland biogeochemistry and ecosystem functions lag behind many mesic systems. To improve our capacity to forecast ecosystem change, we propose focusing on the controls and consequences of two key characteristics affecting dryland biogeochemistry: (1) high spatial and temporal heterogeneity in environmental conditions and (2) generalized resource scarcity. In addition to climate change, drylands are experiencing accelerating land-use change. Building our understanding of dryland biogeochemistry in both intact and disturbed systems will better equip us to address the interacting effects of climate change and landscape degradation. Responding to these challenges will require a diverse, globally distributed and interdisciplinary community of dryland experts united towards better understanding these vast and important ecosystems.

What is a biocrust? A refined, contemporary definition for a broadening research community.

Studies of biological soil crusts (biocrusts) have proliferated over the last few decades. The biocrust literature has broadened, with more studies assessing and describing the function of a variety of biocrust communities in a broad range of biomes and habitats and across a large spectrum of disciplines, and also by the incorporation of biocrusts into global perspectives and biogeochemical models. As the number of biocrust researchers increases, along with the scope of soil communities defined as 'biocrust', it is worth asking whether we all share a clear, universal, and fully articulated definition of what constitutes a biocrust. In this review, we synthesize the literature with the views of new and experienced biocrust researchers, to provide a refined and fully elaborated definition of biocrusts. In doing so, we illustrate the ecological relevance and ecosystem services provided by them. We demonstrate that biocrusts are defined by four distinct elements: physical structure, functional characteristics, habitat, and taxonomic composition. We describe outgroups, which have some, but not all, of the characteristics necessary to be fully consistent with our definition and thus would not be considered biocrusts. We also summarize the wide variety of different types of communities that fall under our definition of biocrusts, in the process of highlighting their global distribution. Finally, we suggest the universal use of the Belnap, Büdel & Lange definition, with minor modifications: Biological soil crusts (biocrusts) result from an intimate association between soil particles and differing proportions of photoautotrophic (e.g. cyanobacteria, algae, lichens, bryophytes) and heterotrophic (e.g. bacteria, fungi, archaea) organisms, which live within, or immediately on top of, the uppermost millimetres of soil. Soil particles are aggregated through the presence and activity of these often extremotolerant biota that desiccate regularly, and the resultant living crust covers the surface of the ground as a coherent layer. With this detailed definition of biocrusts, illustrating their ecological functions and widespread distribution, we hope to stimulate interest in biocrust research and inform various stakeholders (e.g. land managers, land users) on their overall importance to ecosystem and Earth system functioning.

Multiple resource limitation of dryland soil microbial carbon cycling on the Colorado Plateau.

Understanding interactions among biogeochemical cycles is increasingly important as anthropogenic alterations of global climate and of carbon (C), nitrogen (N), and phosphorus (P) cycles interactively affect the Earth system. Ecosystem processes in the dryland biome, which makes up over 40% of Earth's terrestrial surface, are often distinctively sensitive to small changes in resource availability, likely because levels of many resources are low. However, data also suggest that simultaneous changes in the availability of multiple resources may be necessary to affect a response in these low-resource systems, offering an opportunity to test patterns and controls of co-limitation, serial limitation, and individual limitation in soil environments. While drylands may play a governing role in key aspects of Earth's C cycle, and while an improved understanding of resource limitation could substantially improve our forecasts of dryland responses to change, our understanding of interacting controls on soil C cycle processes remains notably poor in these dry systems. Here, we address multiple fundamental hypotheses of resource controls over ecosystem function to test how water, C, N, and P regulate soil C cycling individually and interactively in a dryland ecosystem on the Colorado Plateau. Using a series of laboratory incubations, we found that, while water, C, and N limited C cycling through serial limitation, water alone resulted in an extremely small respiratory response from target organisms, whereas water + C resulted in a dramatic increase in soil C cycling, suggesting a degree of functional co-limitation. Nitrogen additions alone resulted in no changes to soil C cycling, but when N was added in concert with water and C, N greatly increased soil C cycling rates relative to additions of water and C without N. Phosphorus additions had no effect on the C cycle either alone or synergistically. These patterns were consistent with the stoichiometry of the system and interactions among resources were surprising in ways that inform our understanding of critical theories in ecology, such as the Transient Maxima Hypothesis, supporting the suggestion that multiple resource limitation explains pulse-dynamic C cycling in drylands better than water limitation alone.

Incorporating Biogeochemistry into Dryland Restoration.

Dryland degradation is a persistent and accelerating global problem. Although the mechanisms initiating and maintaining dryland degradation are largely understood, returning productivity and function through ecological restoration remains difficult. Water limitation commonly drives slow recovery rates within drylands; however, the altered biogeochemical cycles that accompany degradation also play key roles in limiting restoration outcomes. Addressing biogeochemical changes and resource limitations may help improve restoration efforts within this difficult-to-restore biome. In the present article, we present a synthesis of restoration literature that identifies multiple ways biogeochemical understandings might augment dryland restoration outcomes, including timing restoration around resource cycling and uptake, connecting heterogeneous landscapes, manipulating resource pools, and using organismal functional traits to a restoration advantage. We conclude by suggesting ways to incorporate biogeochemistry into existing restoration frameworks and discuss research directions that may help improve restoration outcomes in the world's highly altered dryland landscapes.

Experimental warming and its legacy effects on root dynamics following two hurricane disturbances in a wet tropical forest.

Tropical forests are expected to experience unprecedented warming and increases in hurricane disturbances in the coming decades; yet, our understanding of how these productive systems, especially their belowground component, will respond to the combined effects of varied environmental changes remains empirically limited. Here we evaluated the responses of root dynamics (production, mortality, and biomass) to soil and understory warming (+4°C) and after two consecutive tropical hurricanes in our in situ warming experiment in a tropical forest of Puerto Rico: Tropical Responses to Altered Climate Experiment (TRACE). We collected minirhizotron images from three warmed plots and three control plots of 12 m2 . Following Hurricanes Irma and María in September 2017, the infrared heater warming treatment was suspended for repairs, which allowed us to explore potential legacy effects of prior warming on forest recovery. We found that warming significantly reduced root production and root biomass over time. Following hurricane disturbance, both root biomass and production increased substantially across all plots; the root biomass increased 2.8-fold in controls but only 1.6-fold in previously warmed plots. This pattern held true for both herbaceous and woody roots, suggesting that the consistent antecedent warming conditions reduced root capacity to recover following hurricane disturbance. Root production and mortality were both related to soil ammonium nitrogen and microbial biomass nitrogen before and after the hurricanes. This experiment has provided an unprecedented look at the complex interactive effects of disturbance and climate change on the root component of a tropical forested ecosystem. A decrease in root production in a warmer world and slower root recovery after a major hurricane disturbance, as observed here, are likely to have longer-term consequences for tropical forest responses to future global change.

Experimental warming across a tropical forest canopy height gradient reveals minimal photosynthetic and respiratory acclimation.

Tropical forest canopies cycle vast amounts of carbon, yet we still have a limited understanding of how these critical ecosystems will respond to climate warming. We implemented in situ leaf-level + 3°C experimental warming from the understory to the upper canopy of two Puerto Rican tropical tree species, Guarea guidonia and Ocotea sintenisii. After approximately 1 month of continuous warming, we assessed adjustments in photosynthesis, chlorophyll fluorescence, stomatal conductance, leaf traits and foliar respiration. Warming did not alter net photosynthetic temperature response for either species; however, the optimum temperature of Ocotea understory leaf photosynthetic electron transport shifted upward. There was no Ocotea respiratory treatment effect, while Guarea respiratory temperature sensitivity (Q10 ) was down-regulated in heated leaves. The optimum temperatures for photosynthesis (Topt ) decreased 3-5°C from understory to the highest canopy position, perhaps due to upper canopy stomatal conductance limitations. Guarea upper canopy Topt was similar to the mean daytime temperatures, while Ocotea canopy leaves often operated above Topt . With minimal acclimation to warmer temperatures in the upper canopy, further warming could put these forests at risk of reduced CO2 uptake, which could weaken the overall carbon sink strength of this tropical forest.

Resistance, Resilience, and Recovery of Dryland Soil Bacterial Communities Across Multiple Disturbances.

Dryland ecosystems are sensitive to perturbations and generally slow to recover post disturbance. The microorganisms residing in dryland soils are especially important as they contribute to soil structure and nutrient cycling. Disturbance can have particularly strong effects on dryland soil structure and function, yet the natural resistance and recovery of the microbial components of dryland soils has not been well documented. In this study, the recovery of surface soil bacterial communities from multiple physical and environmental disturbances is assessed. Samples were collected from three field sites in the vicinity of Moab, UT, United States, 6 to 7 years after physical and climate disturbance manipulations had been terminated, allowing for the assessment of community recovery. Additionally, samples were collected in a transect that included three habitat patches: the canopy zone soils under the dominant shrubs, the interspace soils that are colonized by biological soil crusts, and edge soils at the plot borders. Field site and habitat patch were significant factors structuring the bacterial communities, illustrating that sites and habitats harbored unique soil microbiomes. Across the different sites and disturbance treatments, there was evidence of significant bacterial community recovery, as bacterial biomass and diversity were not significantly different than control plots. There was, however, a small number of 16S rRNA gene amplicon sequence variants that distinguished particular treatments, suggesting that legacy effects of the disturbances still remained. Taken together, these data suggest that dryland bacterial communities may possess a previously unappreciated potential to recover within years of the original disturbance.

Increasing availability of lower energy meals vs. energy labelling in virtual full-service restaurants: two randomized controlled trials in participants of higher and lower socioeconomic position.

BACKGROUND: There are a range of interventions designed to promote healthier food choices in full-service restaurants. However, it is unclear how these interventions affect dietary choices in people of lower and higher socioeconomic position (SEP). METHODS: A total of 2091 US participants recruited online completed Study 1 (n = 1001) and Study 2 (n = 1090). Recruitment was stratified by participant highest education level, resulting in higher SEP and lower SEP groups. In a between-subjects design, participants made hypothetical food choices (main dish, plus optional sides and desserts) from six restaurants menus in the absence vs. presence of menu energy labelling and from menus with baseline (10%) vs. increased availability (50%) of lower energy main dishes. Data were collected and analysed in 2019. Two studies were conducted in order to examine replicability and generalisability of findings across different restaurant menu types. RESULTS: Across both studies, increasing the availability of lower energy main menu options decreased the average energy content of the ordered main dish (- 129 kcal, 95% CI [- 139; - 119]) and total energy ordered (- 117 kcal, 95% CI [- 138; - 95]) in both higher and lower SEP participants. Energy labelling significantly reduced the energy content of ordered main dishes in higher SEP participants (- 41 kcal, 95% CI [- 54; - 29]), but not lower SEP participants (- 5 kcal, 95% CI [- 22; 11]). However, energy labelling reduced total energy ordered (- 83 kcal, 95% CI [- 105; - 60]) irrespective of SEP. CONCLUSIONS: In two virtual experiments, increasing the availability of lower energy restaurant main menu options impacted on main menu dish choice and decreased total energy ordered irrespective of SEP. Energy labelling had a less pronounced effect on total energy ordered and had a larger impact on the energy content of main menu dish choice in higher as opposed to lower SEP participants. TRIAL REGISTRATION: Clinicaltrials.gov NCT04336540 retrospectively registered (7 April, 2020).