Toby M. Maxwell’s research while affiliated with Boise State University and other places

Plants respond to their environment with both short‐term, within‐generation trait plasticity, and long‐term, between‐generation evolutionary changes. However, the relative magnitude of plant responses to short‐ and long‐term changes in the environment remains poorly understood. Shifts in phenological traits can serve as harbingers for responses to environmental change, and both a plant's current and source (i.e., genotype origin) environment can affect plant phenology via plasticity and local adaptation, respectively. To assess the role of current and source environments in explaining variation in flowering phenology of Bromus tectorum, an invasive annual grass, we conducted a replicated common garden experiment using 92 genotypes collected across western North America. Replicates of each genotype were planted in two densities (low = 100 seeds/1 m², high = 100 seeds/0.04 m²) under two different temperature treatments (low = white gravel; high = black gravel; 2.1°C average difference) in a factorial design, replicated across four common garden locations in Idaho and Wyoming, USA. We tested for the effect of current environment (i.e., density treatment, temperature treatment, and common garden location), source environment (i.e., genotype source climate), and their interaction on each plant's flowering phenology. Flowering timing was strongly influenced by a plant's current environment, with plants that experienced warmer current climates and higher densities flowering earlier than those that experienced cooler current climates and lower densities. Genotypes from hot and dry source climates flowered consistently earlier than those from cool and wet source climates, even after accounting for genotype relatedness, suggesting that this genetically based climate cline is a product of natural selection. We found minimal evidence of interactions between current and source environments or genotype‐by‐environment interactions. Phenology was more sensitive to variation in the current climate than to variation in source climate. These results indicate that cheatgrass phenology reflects high levels of plasticity as well as rapid local adaptation. Both processes likely contribute to its current success as a biological invader and its capacity to respond to future environmental change.

Soil organic carbon (‘SOC’) in drylands comprises nearly a third of the global SOC pool and has relatively rapid turnover and thus is a key driver of variability in the global carbon cycle. SOC is also a sensitive indicator of longer-term directional change and disturbance-responses of ecosystem C storage. Biome-scale disruption of the dryland carbon cycle by exotic annual grass invasions (mainly Bromus tectorum, ‘Cheatgrass’) threatens carbon storage and corresponding benefits to soil hydrology and nutrient retention. Past studies on cheatgrass impacts mainly focused on total C, and of the few that evaluated SOC, none compared the very different fractions of SOC, such as relatively unstable particulate organic carbon (POC) or relatively stable, mineral-associated organic carbon (MAOC). We measured SOC and its POC and MAOC constituents in the surface soils of sites that had sagebrush canopies but differed in whether their understories had been invaded by cheatgrass or not, in both warm and relatively colder ecoregions of the western USA. MAOC stocks were 36.1% less in the 0–10 cm depth and 46.1% less in the 10–20 cm depth in the cheatgrass-invaded stands compared to the uninvaded stands of the warmer Colorado Plateau, but not in the cooler and more carbon-rich Wyoming Basin ecoregion. In plots where cheatgrass increased SOC, it was via unstable POC. These findings indicate that cheatgrass effects on the distribution of soil carbon among POC and MAOC fractions may vary among ecoregions, and that cheatgrass can reduce forms of carbon that are otherwise considered stable and ‘secure’, i.e. sequestered.

Ecological disturbance can affect carbon storage and stability and is a key consideration for managing lands to preserve or increase ecosystem carbon to ameliorate the global greenhouse gas problem. Dryland soils are massive carbon reservoirs that are increasingly impacted by species invasions and altered fire regimes, including the exotic-grass-fire cycle in the extensive sagebrush steppe of North America. Direct measurement of total carbon in 1174 samples from landscapes of this region that differed in invasion and wildfire history revealed that their impacts depleted soil carbon by 42–49%, primarily in deep horizons, which could amount to 17.1–20.0 Tg carbon lost across the ~400,000 ha affected annually. Disturbance effects on soil carbon stocks were not synergistic, suggesting that soil carbon was lowered to a floor—i.e. a resistant base-level—beneath which further loss was unlikely. Restoration and maintenance of resilient dryland shrublands/rangelands could stabilize soil carbon at magnitudes relevant to the global carbon cycle.

Local adaptation may facilitate range expansion during invasions, but the mechanisms promoting destructive invasions remain unclear. Cheatgrass ( Bromus tectorum ), native to Eurasia and Africa, has invaded globally, with particularly severe impacts in western North America. We sequenced 307 genotypes and conducted controlled experiments. We found that diverse lineages invaded North America, where long-distance gene flow is common. Ancestry and phenotypic clines in the native range predicted those in the invaded range, indicating pre-adapted genotypes colonized different regions. Common gardens showed directional selection on flowering time that reversed between warm and cold sites, potentially maintaining clines. In the Great Basin, genomic predictions of strong local adaptation identified sites where cheatgrass is most dominant. Preventing new introductions that may fuel adaptation is critical for managing ongoing invasions.

Purpose The sensitivity of wildland plants to temperature can be directly measured using experimental manipulations of temperature in situ. We show that soil surface temperature and plant density (per square meter) have a significant impact on the germination, growth, and phenology of Bromus tectorum L., cheatgrass, a short-statured invasive winter-annual grass, and assess a new experimental temperature manipulation method: the application of black and white gravel to warm and cool the soil surface. Methods We monitored height, seed production, and phenological responses of cheatgrass, seeded into colored gravel at low and high densities at two sites in the western USA: Boise, ID and Cheyenne, WY. Soil surface temperature and volumetric water content were measured to assess treatment effects on soil surface microclimate. Results Black gravel increased mean temperatures of the surface soil by 1.6 and 2.6 °C compared to white gravel in Cheyenne and Boise, respectively, causing 21–24 more days with soil temperatures > 0 °C, earlier cheatgrass germination, and up to 2.8-fold increases in cheatgrass height. Higher seeding density of cheatgrass led to 1.4-fold taller plants on black gravel plots at both sites, but not white gravel at the Boise site, indicating a possible thermal benefit or reduction of water demand due to plant clustering in warmer treatments. Conclusions Manipulating soil-surface albedo altered the soil microclimate and thus growth and phenology of cheatgrass, whose life history and growth form confer a strong dependency on soil-surface conditions.

Annual‐grass invasions are transforming desert ecosystems in ways that affect ecosystem carbon (C) balance, but previous studies do not agree on the pattern, magnitude and direction of changes. A recent meta‐analysis of 41 articles and 386 sites concludes that invasion by annual grasses such as cheatgrass (Bromus tectorum L) reduces C in biomass across the Great Basin (Nagy et al., 2021). Reanalysis reveals that whether cheatgrass affects biomass C stocks is not generalizable, but rather depends on the considerable variation in climate across the subject sites. Our analysis suggests that accurate Great Basin‐scale estimates of cheatgrass effects on C balance are not yet possible. Addition of climate variables to the meta‐analysis reveals that cheatgrass invasion (a) reduced C in above‐ground biomass in relatively summer‐wet sites but not in summer‐dry sites, (b) increased surface soil C in sites with intermediate resistance and resilience classifications (R&R) but not in low R&R sites—that is, mesic/aridic soil climates and (c) did not affect deep soil C. Considering that cheatgrass has expanded most in relatively summer‐dry sites and mesic/aridic sites, omission of climate factors leads to model overestimates of cheatgrass effects on C when extrapolating to larger areas. Estimates of cheatgrass effects on C would also be improved if the analysis considered that (a) perennial grasslands are a common community state in the Great Basin that have intermediary C relative to annual grasslands and sagebrush stands, that is the omission of perennial grasslands from analysis inflates the baseline C storage of uninvaded Great Basin ecosystems, and( b) cheatgrass does not often exist in stable monocultures and soil carbon can reflect current or recent presence of other species. Synthesis and applications. Invasions often reveal heterogeneity in ecosystem structure and function that is not otherwise evident, and the heterogeneity can influence estimation of the net impacts of the invaders. For cheatgrass and other invaders, we propose that formally accounting for the spatial variability of invasion on ecosystem functions will improve the estimation of their net effect on ecosystem C, and thus improve prospects for adjusting management practices to optimize C sequestration.