Unforeseen plant phenotypic diversity in a dry and grazed world.

Earth harbours an extraordinary plant phenotypic diversity1 that is at risk from ongoing global changes2,3. However, it remains unknown how increasing aridity and livestock grazing pressure-two major drivers of global change4-6-shape the trait covariation that underlies plant phenotypic diversity1,7. Here we assessed how covariation among 20 chemical and morphological traits responds to aridity and grazing pressure within global drylands. Our analysis involved 133,769 trait measurements spanning 1,347 observations of 301 perennial plant species surveyed across 326 plots from 6 continents. Crossing an aridity threshold of approximately 0.7 (close to the transition between semi-arid and arid zones) led to an unexpected 88% increase in trait diversity. This threshold appeared in the presence of grazers, and moved toward lower aridity levels with increasing grazing pressure. Moreover, 57% of observed trait diversity occurred only in the most arid and grazed drylands, highlighting the phenotypic uniqueness of these extreme environments. Our work indicates that drylands act as a global reservoir of plant phenotypic diversity and challenge the pervasive view that harsh environmental conditions reduce plant trait diversity8-10. They also highlight that many alternative strategies may enable plants to cope with increases in environmental stress induced by climate change and land-use intensification.

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.

What Could Explain δ13C Signatures in Biocrust Cyanobacteria of Drylands?

Dryland ecosystems are increasing in geographic extent and contribute greatly to interannual variability in global carbon dynamics. Disentangling interactions among dominant primary producers, including plants and autotrophic microbes, can help partition their contributions to dryland C dynamics. We measured the δ13C signatures of biological soil crust cyanobacteria and dominant plant species (C3 and C4) across a regional scale in the southwestern USA to determine if biocrust cyanobacteria were coupled to plant productivity (using plant-derived C mixotrophically), or independent of plant activity (and therefore purely autotrophic). Cyanobacterial assemblages located next to all C3 plants and one C4 species had consistently more negative δ13C (by 2‰) than the cyanobacteria collected from plant interspaces or adjacent to two C4 Bouteloua grass species. The differences among cyanobacterial assemblages in δ13C could not be explained by cyanobacterial community composition, photosynthetic capacity, or any measured leaf or root characteristics (all slopes not different from zero). Thus, microsite differences in abiotic conditions near plants, rather than biotic interactions, remain a likely mechanism underlying the observed δ13C patterns to be tested experimentally.

Hotspots of biogeochemical activity linked to aridity and plant traits across global drylands.

Perennial plants create productive and biodiverse hotspots, known as fertile islands, beneath their canopies. These hotspots largely determine the structure and functioning of drylands worldwide. Despite their ubiquity, the factors controlling fertile islands under conditions of contrasting grazing by livestock, the most prevalent land use in drylands, remain virtually unknown. Here we evaluated the relative importance of grazing pressure and herbivore type, climate and plant functional traits on 24 soil physical and chemical attributes that represent proxies of key ecosystem services related to decomposition, soil fertility, and soil and water conservation. To do this, we conducted a standardized global survey of 288 plots at 88 sites in 25 countries worldwide. We show that aridity and plant traits are the major factors associated with the magnitude of plant effects on fertile islands in grazed drylands worldwide. Grazing pressure had little influence on the capacity of plants to support fertile islands. Taller and wider shrubs and grasses supported stronger island effects. Stable and functional soils tended to be linked to species-rich sites with taller plants. Together, our findings dispel the notion that grazing pressure or herbivore type are linked to the formation or intensification of fertile islands in drylands. Rather, our study suggests that changes in aridity, and processes that alter island identity and therefore plant traits, will have marked effects on how perennial plants support and maintain the functioning of drylands in a more arid and grazed world.

Biocrust carbon exchange varies with crust type and time on Chihuahuan Desert gypsum soils.

Introduction: In dryland systems, biological soil crusts (biocrusts) can occupy large areas of plant interspaces, where they fix carbon following rain. Although distinct biocrust types contain different dominant photoautotrophs, few studies to date have documented carbon exchange over time from various biocrust types. This is especially true for gypsum soils. Our objective was to assess the carbon exchange of biocrust types established at the world's largest gypsum dune field at White Sands National Park. Methods: We sampled five different biocrust types from a sand sheet location in three different years and seasons (summer 2020, fall 2021, and winter 2022) for carbon exchange measurements in controlled lab conditions. Biocrusts were rehydrated to full saturation and light incubated for 30 min, 2, 6, 12, 24, and 36 h. Samples were then subject to a 12-point light regime with a LI-6400XT photosynthesis system to determine carbon exchange. Results: Biocrust carbon exchange values differed by biocrust type, by incubation time since wetting, and by date of field sampling. Lichens and mosses had higher gross and net carbon fixation rates than dark and light cyanobacterial crusts. High respiration rates were found after 0.5 h and 2 h incubation times as communities recovered from desiccation, leveling off after 6 h incubation. Net carbon fixation of all types increased with longer incubation time, primarily as a result of decreasing respiration, which suggests rapid recovery of biocrust photosynthesis across types. However, net carbon fixation rates varied from year to year, likely as a product of time since the last rain event and environmental conditions preceding collection, with moss crusts being most sensitive to environmental stress at our study sites. Discussion: Given the complexity of patterns discovered in our study, it is especially important to consider a multitude of factors when comparing biocrust carbon exchange rates across studies. Understanding the dynamics of biocrust carbon fixation in distinct crust types will enable greater precision of carbon cycling models and improved forecasting of impacts of global climate change on dryland carbon cycling and ecosystem functioning.

Disturbance to biocrusts decreased cyanobacteria, N-fixer abundance, and grass leaf N but increased fungal abundance.

Interactions between plants and soil microbes influence plant nutrient transformations, including nitrogen (N) fixation, nutrient mineralization, and resource exchanges through fungal networks. Physical disturbances to soils can disrupt soil microbes and associated processes that support plant and microbial productivity. In low resource drylands, biological soil crusts ("biocrusts") occupy surface soils and house key autotrophic and diazotrophic bacteria, non-vascular plants, or lichens. Interactions among biocrusts, plants, and fungal networks between them are hypothesized to drive carbon and nutrient dynamics; however, comparisons across ecosystems are needed to generalize how soil disturbances alter microbial communities and their contributions to N pools and transformations. To evaluate linkages among plants, fungi, and biocrusts, we disturbed all unvegetated surfaces with human foot trampling twice yearly from 2013-2019 in dry conditions in cyanobacteria-dominated biocrusts in the Chihuahuan Desert grassland and shrubland ecosystems. After 5 years, disturbance decreased the abundances of cyanobacteria (especially Microcoleus steenstrupii clade) and N-fixers (Scytonema sp., and Schizothrix sp.) by >77% and chlorophyll a by up to 55% but, conversely, increased soil fungal abundance by 50% compared with controls. Responses of root-associated fungi differed between the two dominant plant species and ecosystem types, with a maximum of 80% more aseptate hyphae in disturbed than in control plots. Although disturbance did not affect 15 N tracer transfer from biocrusts to the dominant grass, Bouteloua eriopoda, disturbance increased available soil N by 65% in the shrubland, and decreased leaf N of B. eriopoda by up to 16%, suggesting that, although rapid N transfer during peak production was not affected by disturbance, over the long-term plant nutrient content was disrupted. Altogether, the shrubland may be more resilient to detrimental changes due to disturbance than grassland, and these results demonstrated that disturbances to soil microbial communities have the potential to cause substantial changes in N pools by reducing and reordering biocrust taxa.

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.

Grazing and ecosystem service delivery in global drylands.

Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure.

Concurrent and lagged impacts of an anomalously warm year on autotrophic and heterotrophic components of soil respiration: a deconvolution analysis.

*Partitioning soil respiration into autotrophic (R(A)) and heterotrophic (R(H)) components is critical for understanding their differential responses to climate warming. *Here, we used a deconvolution analysis to partition soil respiration in a pulse warming experiment. We first conducted a sensitivity analysis to determine which parameters can be identified by soil respiration data. A Markov chain Monte Carlo technique was then used to optimize those identifiable parameters in a terrestrial ecosystem model. Finally, the optimized parameters were employed to quantify R(A) and R(H) in a forward analysis. *Our results displayed that more than one-half of parameters were constrained by daily soil respiration data. The optimized model simulation showed that warming stimulated R(H) and had little effect on R(A) in the first 2 months, but decreased both R(H) and R(A) during the remainder of the treatment and post-treatment years. Clipping of above-ground biomass stimulated the warming effect on R(H) but not on R(A). Overall, warming decreased R(A) and R(H) significantly, by 28.9% and 24.9%, respectively, during the treatment year and by 27.3% and 33.3%, respectively, during the post-treatment year, largely as a result of decreased canopy greenness and biomass. *Lagged effects of climate anomalies on soil respiration and its components are important in assessing terrestrial carbon cycle feedbacks to climate warming.

Limited effects of early snowmelt on plants, decomposers, and soil nutrients in Arctic tundra soils.

In addition to warming temperatures, Arctic ecosystems are responding to climate change with earlier snowmelt and soil thaw. Earlier snowmelt has been examined infrequently in field experiments, and we lack a comprehensive look at belowground responses of the soil biogeochemical system that includes plant roots, decomposers, and soil nutrients. We experimentally advanced the timing of snowmelt in factorial combination with an open-top chamber warming treatment over a 3-year period and evaluated the responses of decomposers and nutrient cycling processes. We tested two alternative hypotheses: (a) Early snowmelt and warming advance the timing of root growth and nutrient uptake, altering the timing of microbial and invertebrate activity and key nutrient cycling events; and (b) loss of insulating snow cover damages plants, leading to reductions in root growth and altered biological activity. During the 3 years of our study (2010-2012), we advanced snowmelt by 4, 15, and 10 days, respectively. Despite advancing aboveground plant phenology, particularly in the year with the warmest early-season temperatures (2012), belowground effects were primarily seen only on the first sampling date of the season or restricted to particular years or soil type. Overall, consistent and substantial responses to early snowmelt were not observed, counter to both of our hypotheses. The data on soil physical conditions, as well interannual comparisons of our results, suggest that this limited response was because of the earlier date of snowmelt that did not coincide with substantially warmer air and soil temperatures as they might in response to a natural climate event. We conclude that the interaction of snowmelt timing with soil temperatures is important to how the ecosystem will respond, but that 1- to 2-week changes in timing of snowmelt alone are not enough to drive season-long changes in soil microbial and nutrient cycling processes.

Effects of the exposure of TiO2 nanoparticles on basil (Ocimum basilicum) for two generations.

There is a lack of information about the transgenerational effects of titanium dioxide nanoparticles (nano-TiO2) in plants. This study aimed to evaluate the impacts of successive exposure of nano-TiO2 with different surface properties to basil (Ocimum basilicum). Seeds from plants exposed or re-exposed to pristine, hydrophobic, or hydrophilic nano-TiO2 were cultivated for 65 days in soil unamended or amended with 750 mg·kg-1 of the respective particles. Plant growth, concentration of titanium and essential elements, as well as content of carbohydrates and chlorophyll were evaluated. There were no differences on Ti concentration in roots of plants sequentially exposed to pristine or hydrophobic nano-TiO2, or in roots of plants exposed to the corresponding particle, only in the second cycle. However, sequential exposure to hydrophilic particles resulted in 65.2% less Ti in roots, compared to roots of plants exposed the same particles, only in the second cycle. The Ti concentrations in shoots were similar in all treatments. On the other hand, pristine and hydrophilic particles reduced Mg in root by 115% and 81%, respectively, while pristine and hydrophobic particles reduced Ni in shoot by 84% and 75%, respectively, compared to unexposed plants in both cycles. Sequential exposure to pristine nano-TiO2 increased stomatal conductance (214%, p ≤ 0.10), compared to plants that were never exposed. Hydrophobic and hydrophilic nano-TiO2 reduced chlorophyll b (52%) and total chlorophyll (30%) but increased total sugar (186%) and reducing sugar (145%), compared to unexposed plants in both cycles. Sequential exposure to hydrophobic or hydrophilic nano-TiO2 resulted in more adverse effects on photosynthesis but in positive effects on plant growth, compared to pristine nano-TiO2.

Earlier snowmelt and warming lead to earlier but not necessarily more plant growth.

Climate change over the past ∼50 years has resulted in earlier occurrence of plant life-cycle events for many species. Across temperate, boreal and polar latitudes, earlier seasonal warming is considered the key mechanism leading to earlier leaf expansion and growth. Yet, in seasonally snow-covered ecosystems, the timing of spring plant growth may also be cued by snowmelt, which may occur earlier in a warmer climate. Multiple environmental cues protect plants from growing too early, but to understand how climate change will alter the timing and magnitude of plant growth, experiments need to independently manipulate temperature and snowmelt. Here, we demonstrate that altered seasonality through experimental warming and earlier snowmelt led to earlier plant growth, but the aboveground production response varied among plant functional groups. Earlier snowmelt without warming led to early leaf emergence, but often slowed the rate of leaf expansion and had limited effects on aboveground production. Experimental warming alone had small and inconsistent effects on aboveground phenology, while the effect of the combined treatment resembled that of early snowmelt alone. Experimental warming led to greater aboveground production among the graminoids, limited changes among deciduous shrubs and decreased production in one of the dominant evergreen shrubs. As a result, we predict that early onset of the growing season may favour early growing plant species, even those that do not shift the timing of leaf expansion.