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Observations of net soil exchange of CO 2 in a dryland show experimental warming increases carbon losses in biocrust soils

Many arid and semiarid ecosystems have soils covered with well-developed biological soil crust communities (biocrusts) made up of mosses, lichens, cyanobacteria, and heterotrophs living at the soil surface. These communities are a fundamental component of dryland ecosystems, and are critical to dryland carbon (C) cycling. To examine the effects of warming temperatures on soil C balance in a dryland ecosystem, we used infrared heaters to warm biocrust-dominated soils to 2 °C above control conditions at a field site on the Colorado Plateau, USA. We monitored net soil exchange (NSE) of CO 2 every hour for 21 months using automated flux chambers (5 control and 5 warmed chambers), which included the CO 2 fluxes of the biocrusts and the soil beneath them. We observed measurable photosynthesis in biocrust soils on 12 % of measurement days, which correlated well with precipitation events and soil wet-up. These days included several snow events, providing what we believe to be the first evidence of substantial photosynthesis underneath snow by biocrust organisms in drylands. Overall, biocrust soils in both control and warmed plots were net CO 2 sources to the atmosphere, with control plots losing 62 ± 8 g C m −2 (mean ± SE) over the first year of measurement and warmed plots losing 74 ± 9 g C m −2 . Between control and warmed plots, the difference in soil C loss was uncertain over the course of the entire year due to large and variable rates in spring, but on days during which soils were wet and crusts were actively photosynthesizing, biocrusts that were warmed by 2 °C had a substantially more negative C balance (i.e., biocrust soils took up less C and/or lost more C in warmed plots). Taken together, our data suggest a substantial risk of increased C loss from biocrust soils with higher future temperatures, and highlight a robust capacity to predict CO 2 exchange in biocrust soils using easily measured environmental parameters.

Arizona, Colorado, New Mexico, Utah

Unraveling mechnisms underlying effects of wetting–drying cycles on soil respiration in a dryland

Rewetting of dry soils usually stimulates soil carbon (C) emission, a phenomenon known as the Birch effect. Soil C cycling in drylands, which store approximately one third of terrestrial soil organic C (SOC), is strongly affected by wetting–drying cycles. However, the physical, chemical, and biological mechanisms that link rewetting cycles with dryland soil C cycling have not been comprehensively studied, nor do we understand how these mechanisms interact with each other. Here, we conducted a dryland soil incubation experiment manipulating four factors related to global change (soil moisture content, soil moisture variability, C availability, and prior warming) in a factorial design. The experiment was divided into two periods: a rewetting period consisting of six 14-d wetting–drying cycles; and a recovery period lasting 28 days during which soil moisture content was held stable, allowing for examination of the legacy effects of the wet-dry cycles. Rewetting cycles decreased soil aggregate stability under some conditions, but their effects on soil microbial biomass and fungal communities, soil enzyme activities, soil priming, and soil dissolved C were not significant. We found lower average soil respiration under the wetting–drying treatment than the stable soil moisture treatment, and Birch effects were observed, but only under some conditions. This was probably because moisture variability exacerbated soil microbial metabolic stress, which showed itself as oxygen limitation during the initial precipitation pulse and as water limitation during soil drying. Notably, respiration rates remained low even after moisture fluctuations ceased, suggesting a legacy effect of rewetting cycles on dryland microbial communities. Overall, rewetting inhibited aggregate formation (physical mechanism), and suppressed soil respiration by inducing soil microbial metabolic stress (biological mechanism), ultimately leading to lower soil C loss under rewetting. Our findings indicate that Birch effects are mediated by the magnitude of moisture variability, the availability of C, and the degree of physiological stress microbes experience.

Biogeochemistry

Performance of the ecosystem demography model (EDv2.2) in simulating gross primary production capacity and activity in a dryland study area

Dryland ecosystems play an important role in the global carbon cycle, including regulating the inter-annual global carbon sink. Dynamic global vegetation models (DGVMs) are essential tools that can help us better understand carbon cycling in different ecosystems. Currently, there is limited knowledge of the performance of these models in drylands partly due to characterizing the heterogeneity of the vegetation and hydrometeorological conditions. The aim of this study is to evaluate the performance of a DGVM for drylands to facilitate improved understanding of gross primary production (GPP) as one of the important components of the carbon cycle. We performed a sensitivity analysis and calibrated the Ecosystem Demography (EDv2.2) DGVM to simulate GPP in a dryland watershed (Reynolds Creek Experimental Watershed, Idaho) in the western US for the years 2000-2017. GPP capacity and activity were investigated by comparing model simulations with GPP estimated from eddy covariance data (available from 2015-2017) and remote sensing products (2000-2017). Our results show good performance of EDv2.2 at daily timesteps ( R M S E ≈ 0.38 [ kgC / m 2 / year ] ) "> RMSE≈0.38[kgC/m2/year])

Idaho

Africa’s drylands in a changing world: Challenges for wildlife conservation under climate and land-use changes in the Greater Etosha Landscape

Proclaimed in 1907, Etosha National Park in northern Namibia is an iconic dryland system with a rich history of wildlife conservation and research. A recent research symposium on wildlife conservation in the Greater Etosha Landscape (GEL) highlighted increased concern of how intensification of global change will affect wildlife conservation based on participant responses to a questionnaire. The GEL includes Etosha and surrounding areas, the latter divided by a veterinary fence into large, private farms to the south and communal areas of residential and farming land to the north. Here, we leverage our knowledge of this ecosystem to provide insight into the broader challenges facing wildlife conservation in this vulnerable dryland environment. We first look backward, summarizing the history of wildlife conservation and research trends in the GEL based on a literature review, providing a broad-scale understanding of the socioecological processes that drive dryland system dynamics. We then look forward, focusing on eight key areas of challenge and opportunity for this ecosystem: climate change, water availability and quality, vegetation and fire management, adaptability of wildlife populations, disease risk, human-wildlife conflict, wildlife crime, and human dimensions of wildlife conservation. Using this model system, we summarize key lessons and identify critical threats highlighting future research needs to support wildlife management. Research in the GEL has followed a trajectory seen elsewhere reflecting an increase in complexity and integration across biological scales over time. Yet, despite these trends, a gap exists between the scope of recent research efforts and the needs of wildlife conservation to adapt to climate and land-use changes. Given the complex nature of climate change, in addition to locally existing system stressors, a framework of forward-thinking adaptive management to address these challenges, supported by integrative and multidisciplinary research could be beneficial. One critical area for growth is to better integrate research and wildlife management across land-use types. Such efforts have the potential to support wildlife conservation efforts and human development goals, while building resilience against the impacts of climate change. While our conclusions reflect the specifics of the GEL ecosystem, they have direct relevance for other African dryland systems impacted by global change.

Africa, Greater Etosha Landscape

Dryland mechanisms could widely control ecosystem functioning in a drier and warmer world

Responses of terrestrial ecosystems to climate change have been explored in many regions worldwide. While continued drying and warming may alter process rates and deteriorate the state and performance of ecosystems, it could also lead to more fundamental changes in the mechanisms governing ecosystem functioning. Here we argue that climate change will induce unprecedented shifts in these mechanisms in historically wetter climatic zones, towards mechanisms currently prevalent in dry regions, which we refer to as ‘dryland mechanisms’. We discuss 12 dryland mechanisms affecting multiple processes of ecosystem functioning, including vegetation development, water flow, energy budget, carbon and nutrient cycling, plant production and organic matter decomposition. We then examine mostly rare examples of the operation of these mechanisms in non-dryland regions where they have been considered irrelevant at present. Current and future climate trends could force microclimatic conditions across thresholds and lead to the emergence of dryland mechanisms and their increasing control over ecosystem functioning in many biomes on Earth.

Nature Ecology & Evolution

Abiotic and biotic controls of non-native perennial plant success in drylands

Drivers of non-native plant success in drylands are poorly understood. Here we identify functional differences between dryland native and non-native perennial plants and assess how biotic, abiotic and anthropogenic factors shape the success of the latter. On the basis of plant community and functional trait data from 98 sites across 25 countries, we report a total of 41 non-native plant species at 31 sites. Non-natives tend towards faster growth strategies than natives. Non-native plant richness is higher at sites with greater grazing pressure and under environmental conditions associated with higher soil fertility, decomposition and fungal richness—conditions that tend to occur in less arid regions—and lower where native plant and herbivore richness are greater. Non-native plant cover correlates positively with grazing pressure and negatively with native plant richness. Taken together, our results suggest that non-native plant success in drylands is facilitated when high grazing pressure coincides with elevated resource availability. Such context-dependence of non-native plant success and linkages with native plant and herbivore diversity highlight the need for managing grazing and conserving biodiversity across the world’s drylands.

Nature Ecology and Evolution

Connectivity dynamics in dryland litter cycles: Moving decomposition beyond spatial stasis

Drylands (arid and semiarid ecosystems) cover nearly half of Earth's terrestrial surface, but biogeochemical pools and processes in these systems remain poorly understood. Litter can account for a substantial portion of carbon and nutrient pools in these systems, with litter decomposition exerting important controls over biogeochemical cycling. Dryland decomposition is typically treated as a spatially static process in which litter is retained and decomposed where it is initially deposited. Although this assumption is reasonable for mesic systems with continuous plant canopy cover and a stable subcanopy litter layer, dryland pools generally reflect discontinuous inputs from heterogeneous canopy cover followed by substantial litter transport. In the present article, we review horizontal and vertical transport processes that move litter from the initial deposition point and retention elements that influence litter accumulation patterns. Appreciation of the spatially dynamic litter cycle, including quantitative assessment of transport patterns, will improve estimates of the fate and distribution of organic matter in current and future drylands.

BioScience

Does restoration of plant diversity trigger concomitant soil microbiome changes in dryland ecosystems?

Drylands are highly vulnerable to land degradation, and despite increasing efforts, restoration success remains low. Although often ignored in the design and deployment of management strategies, soil microbial communities might be critical for dryland restoration due to their central role in promoting soil stability, nutrient cycling and plant establishment. We collected soil samples from eight dryland restoration sites within RestoreNet, a restoration field trial network, and determined their soil microbiome using 16S rRNA (bacteria and archaea) and ITS (fungi) amplicon sequencing. Each previously degraded site was treated with monoculture (single species) and polyculture (multiple species) seedling plantings. Contrary to our initial expectations, we found that these different revegetation interventions did not trigger changes in microbial diversity, composition or relative abundance of functional groups across sites after 1 year of revegetation. Synthesis and applications . Considering the crucial role of soil micro-organisms in dryland ecosystem functions, our results suggest that site-specific targeted microbiome restoration should be considered to accelerate the establishment of desired microbial communities. Plant community-based restoration practices such as revegetation have a limited impact on soil micro-organisms in the short term.

Journal of Applied Ecology

Plant water-use strategies predict restoration success across degraded drylands

Plant strategies for coping with water limitation are likely to mediate restoration outcomes in degraded dryland ecosystems. Trade-offs in traits related to water acquisition and use can intensify in more arid environments, making their effects on dryland restoration success even more salient. However, isolating the effects of drought responses from those of other environmental factors, as well as identifying the specific drought resistance traits that influence restoration success, can be difficult. In the present study, we couple a controlled dry-down experiment with a cross-site restoration field trial of out-planted seedlings (RestoreNet) using a suite of dryland herbaceous plant species from the same seed sources. We quantified interspecific variation in physiological responses to drought, specifically reductions in stomatal conductance ( g s ) and stem water potential (SWP), by comparing well-watered control plants to those experiencing decreasing soil moisture. Drought responses of SWP and g s varied independently among species, but both were related to survival in the cross-site restoration field trial when effect sizes were aggregated across all sites. Responses were consistent with acquisitive water-use strategies resulting in greater success, where species with greater declines in SWP or weaker declines in g s under drought had greater survival. The correlation between SWP drought response and survival also intensified in sites with lower accumulated precipitation following restoration. Differences among functional groups revealed two different paths to restoration success: forbs that maintain high g s and narrow safety margins to maximize exploitation of moisture pulses before going into drought dormancy, or C 4 grasses that maintain efficient water uptake in drying soils while risking cavitation. C 3 grass species varied between these two strategies. Synthesis and applications . Taken together, the results of this study and others conducted at RestoreNet sites indicate that while a diversity of physiological responses to drought may exist in dryland plant communities, successfully restoring herbaceous species through out-planting in degraded conditions is likely to be achieved with species that maximize water uptake via one of two strategies, with tolerance of low SWP being particularly important in the most arid settings.

Journal of Applied Ecology

Biotic vs abiotic controls on temporal sensitivity of primary production to precipitation across North American drylands

Dryland net primary productivity (NPP) is sensitive to temporal variation in precipitation (PPT), but the magnitude of this ‘temporal sensitivity’ varies spatially. Hypotheses for spatial variation in temporal sensitivity have often emphasized abiotic factors, such as moisture limitation, while overlooking biotic factors, such as vegetation structure. We tested these hypotheses using spatiotemporal models fit to remote-sensing data sets to assess how vegetation structure and climate influence temporal sensitivity across five dryland ecoregions of the western USA. Temporal sensitivity was higher in locations and ecoregions dominated by herbaceous vegetation. By contrast, much less spatial variation in temporal sensitivity was explained by mean annual PPT. In fact, ecoregion-specific models showed inconsistent associations of sensitivity and PPT; whereas sensitivity decreased with increasing mean annual PPT in most ecoregions, it increased with mean annual PPT in the most arid ecoregion, the hot deserts. The strong, positive influence of herbaceous vegetation on temporal sensitivity indicates that herbaceous-dominated drylands will be particularly sensitive to future increases in precipitation variability and that dramatic changes in cover type caused by invasions or shrub encroachment will lead to changes in dryland NPP dynamics, perhaps independent of changes in precipitation.

New Phytologist

Spatially explicit patterns in a dryland's soil respiration and relationships with climate, whole plant photosynthesis and soil fertility

Arid and semiarid ecosystems play a significant role in regulating global carbon cycling, yet our understanding of the controls over the dominant pathways of dryland CO 2 exchange remains poor. Substantial amounts of dryland soil are not covered by vascular plants and this patchiness in cover has important implications for spatial patterns and controls of carbon cycling. Spatial variation in soil respiration has been attributed to variation in soil moisture, temperature, nutrients and rhizodeposition, while seasonal patterns have been attributed to changes in moisture, temperature and photosynthetic inputs belowground. To characterize how controls over respiration vary spatially and temporally in a dryland ecosystem and to concurrently explore multiple potential controls, we estimated whole plant net photosynthesis (A net ) and soil respiration at four distances from the plant base, as well as corresponding fine root biomass and soil carbon and nitrogen pools, four times during a growing season. To determine if the controls vary between different plant functional types for Colorado Plateau species, measurements were made on the C 4 shrub, Atriplex confertifolia , and C 3 grass, Achnatherum hymenoides . Soil respiration declined throughout the growing season and diminished with distance from the plant base, though variations in both were much smaller than expected. The strongest relationship was between soil respiration and soil moisture. Soil respiration was correlated with whole plant A net , although the relationship varied between species and distance from plant base. In the especially dry year of this study we did not observe any consistent correlations between soil respiration and soil carbon or nitrogen pools. Our findings suggest that abiotic factors, especially soil moisture, strongly regulate the response of soil respiration to biotic factors and soil carbon and nitrogen pools in dryland communities and, at least in dry years, may override expected spatial and seasonal patterns.

Oikos

Disturbance amplifies sensitivity of dryland productivity to precipitation variability

Variability of the terrestrial global carbon sink is largely determined by the response of dryland productivity to annual precipitation. Despite extensive disturbance in drylands, how disturbance alters productivity-precipitation relationships remains poorly understood. Using remote-sensing to pair more than 5600 km of natural gas pipeline corridors with neighboring undisturbed areas in North American drylands, we found that disturbance reduced average annual production 6 to 29% and caused up to a fivefold increase in the sensitivity of net primary productivity (NPP) to interannual variation in precipitation. Disturbance impacts were larger and longer-lasting at locations with higher precipitation (>450 mm mean annual precipitation). Disturbance effects on NPP dynamics were mostly explained by shifts from woody to herbaceous vegetation. Severe disturbance will amplify effects of increasing precipitation variability on NPP in drylands.

Science Advances

Drivers of woody dominance across global drylands

Increases in the abundance of woody species have been reported to affect the provisioning of ecosystem services in drylands worldwide. However, it is virtually unknown how multiple biotic and abiotic drivers, such as climate, grazing, and fire, interact to determine woody dominance across global drylands. We conducted a standardized field survey in 304 plots across 25 countries to assess how climatic features, soil properties, grazing, and fire affect woody dominance in dryland rangelands. Precipitation, temperature, and grazing were key determinants of tree and shrub dominance. The effects of grazing were determined not solely by grazing pressure but also by the dominant livestock species. Interactions between soil, climate, and grazing and differences in responses to these factors between trees and shrubs were key to understanding changes in woody dominance. Our findings suggest that projected changes in climate and grazing pressure may increase woody dominance in drylands, altering their structure and functioning.

Science Advances

Restoration of Native Hawaiian Dryland Forest at Auwahi, Maui

BACKGROUND The powerful volcanoes that formed the high islands of the Hawaiian archipelago block northeasterly tradewinds, creating wet, windward rain forests and much drier, leeward forests. Dryland forests in Hawai'i receive only about 20 inches of rain a year. However, the trees in these forests intercept fog and increase ground moisture levels, thereby enabling these seemingly inhospitable habitats to support a diverse assemblage of plants and animals. Dryland forests of the Hawaiian Islands, like those worldwide, have been heavily impacted by humans both directly and indirectly. Less than 10% of Hawai'i's original dryland forest habitat remains. These forests have been severely impacted by urban development, ranching and agriculture, and invasive species. In particular, browsing animals and alien grasses have caused significant damage. Feral ungulates, including goats, sheep, cattle, and pigs, consume sensitive plants. Alien grasses have become dominant in the understory in many dryland habitats. In addition, these introduced grasses are fire-adapted and have increased the incidence of wildfire in these ecosystems. Native Hawaiian plants did not evolve with frequent fires or mammalian herbivores and typically do not survive well under these pressures.

Fact Sheet

Understanding the role of ecohydrological feedbacks in ecosystem state change in drylands

Ecohydrological feedbacks are likely to be critical for understanding the mechanisms by which changes in exogenous forces result in ecosystem state change. We propose that in drylands, the dynamics of ecosystem state change are determined by changes in the type (stabilizing vs amplifying) and strength of ecohydrological feedbacks following a change in exogenous forces. Using a selection of five case studies from drylands, we explore the characteristics of ecohydrological feedbacks and resulting dynamics of ecosystem state change. We surmise that stabilizing feedbacks are critical for the provision of plant-essential resources in drylands. Exogenous forces that break these stabilizing feedbacks can alter the state of the system, although such changes are potentially reversible if strong amplifying ecohydrological feedbacks do not develop. The case studies indicate that if amplifying ecohydrological feedbacks do develop, they are typically associated with abiotic processes such as runoff, erosion (by wind and water), and fire. These amplifying ecohydrological feedbacks progressively modify the system in ways that are long-lasting and possibly irreversible on human timescales.

Ecohydrology

Climate change may restrict dryland forest regeneration in the 21st century

The persistence and geographic expansion of dryland forests in the 21st century will be influenced by how climate change supports the demographic processes associated with tree regeneration. Yet, the way that climate change may alter regeneration is unclear. We developed a quantitative framework that estimates forest regeneration potential (RP) as a function of key environmental conditions for ponderosa pine, a key dryland forest species. We integrated meteorological data and climate projections for 47 ponderosa pine forest sites across the western United States, and evaluated RP using an ecosystem water balance model. Our primary goal was to contrast conditions supporting regeneration among historical, mid-21st century and late-21st century time frames. Future climatic conditions supported 50% higher RP in 2020–2059 relative to 1910–2014. As temperatures increased more substantially in 2060–2099, seedling survival decreased, RP declined by 50%, and the frequency of years with very low RP increased from 25% to 58%. Thus, climate change may initially support higher RP and increase the likelihood of successful regeneration events, yet will ultimately reduce average RP and the frequency of years with moderate climate support of regeneration. Our results suggest that climate change alone may begin to restrict the persistence and expansion of dryland forests by limiting seedling survival in the late 21st century.

Ecology

Biological soil crusts as an organizing principle in drylands

Biological soil crusts (biocrusts) have been present on Earth’s terrestrial surfaces for billions of years. They are a critical part of ecosystem processes in dryland regions, as they cover most of the soil surface and thus mediate almost all inputs and outputs from soils in these areas. There are many intriguing, but understudied, roles these communities may play in drylands. These include their function in nutrient capture and transformation, influence on the movement and distribution of nutrients and water within dryland soils, ability to structure vascular plant communities, role in creating biodiversity hotspots, and the possibility that they can be used as indicators of soil health. There are still many fascinating aspects of these communities that need study, and we hope that this chapter will facilitate such efforts.

Book chapter

Biological phosphorus cycling in dryland regions

The relatively few studies done on phosphorus (P) cycling in arid and semiarid lands (drylands) show many factors that distinguish P cycling in drylands from that in more mesic regions. In drylands, most biologically relevant P inputs and losses are from the deposition and loss of dust. Horizontal and vertical redistribution of P is an important process. P is concentrated at the soil surface and thus vulnerable to loss via erosion. High pH and CaCO 3 limit P bioavailability, and low rainfall limits microbe and plant ability to free abiotically bound P via exudates, thus making it available for uptake. Many invasive plants are able to access recalcitrant P more effectively than are native plants. As P availability depends on soil moisture and temperature, climate change is expected to have large impacts on P cycling.

Book chapter