Phenology of perennial, native grass, belowground axillary buds in the northern mixed-grass prairie.

PREMISE OF THE STUDY: Vegetative reproduction from belowground bud banks is the primary driver of grassland systems. Despite the importance of bud banks, the timing of recruitment and the crucial link between formation and maintenance is unknown. METHODS: We assessed patterns of belowground bud development, dormancy, and mortality associated with three perennial native grasses in the northern Great Plains. Temperature and soil moisture were measured below the soil surface to determine relationships with belowground bud development. KEY RESULTS: Blue grama (Bouteloua gracilis) generated more buds over winter that remained dormant; whereas, C3 species needle-and-thread (Hesperostipa comata) and western wheatgrass (Pascopyrum smithii), maintained limited dormant buds throughout winter. Soil temperature was a good predictor for C4 species bud production; whereas, soil moisture was a reliable predictor for C3 buds. Distinct differences existed between C4 species blue grama and C3 species needle-and-thread, whereas C3 species western wheatgrass (Pascopyrum smithii) was intermediate, indicating there is likely a species-specific continuum between the C3 and C4 extremes rather than a stark difference. CONCLUSIONS: The ability to predict belowground bud development is a novel insight to native perennial grasses. Native grass species' strategies and adaptability regarding belowground bud bank size and bud phenology are important factors optimizing tiller recruitment given the variable growing conditions. Patterns of bud dormancy and development will provide insight to the underlying mechanisms by which management practices and fluctuations in precipitation amount and growing season length can alter mixed-grass prairie plant community dynamics.

Climatic controls of aboveground net primary production in semi-arid grasslands along a latitudinal gradient portend low sensitivity to warming.

Although climate models forecast warmer temperatures with a high degree of certainty, precipitation is the primary driver of aboveground net primary production (ANPP) in most grasslands. Conversely, variations in temperature seldom are related to patterns of ANPP. Thus forecasting responses to warming is a challenge, and raises the question: how sensitive will grassland ANPP be to warming? We evaluated climate and multi-year ANPP data (67 years) from eight western US grasslands arrayed along mean annual temperature (MAT; ~7-14 °C) and mean annual precipitation (MAP; ~250-500 mm) gradients. We used regression and analysis of covariance to assess relationships between ANPP and temperature, as well as precipitation (annual and growing season) to evaluate temperature sensitivity of ANPP. We also related ANPP to the standardized precipitation evaporation index (SPEI), which combines precipitation and evapotranspiration to better represent moisture available for plant growth. Regression models indicated that variation in growing season temperature was negatively related to total and graminoid ANPP, but precipitation was a stronger predictor than temperature. Growing season temperature was also a significant parameter in more complex models, but again precipitation was consistently a stronger predictor of ANPP. Surprisingly, neither annual nor growing season SPEI were as strongly related to ANPP as precipitation. We conclude that forecasted warming likely will affect ANPP in these grasslands, but that predicting temperature effects from natural climatic gradients is difficult. This is because, unlike precipitation, warming effects can be positive or negative and moderated by shifts in the C3/C4 ratios of plant communities.

Omitted variable bias in studies of plant interactions.

Models of plant-plant interactions underpin our understanding of species coexistence, invasive plant impacts, and plant community responses to climate change. In recent studies, models of competitive interactions failed predictive tests, thereby casting doubt on results of many past studies. We believe these model failures owe at least partly to heterogeneity in unmodeled factors (e.g., nutrients, soil pathogens) that affect both target plants and neighboring competitors. Such heterogeneity is ubiquitous, and models that do not account for it will suffer omitted variable bias. We used instrumental variables analysis to test for and correct omitted variable bias in studies that followed common protocols for measuring plant competition. In an observational study, omitted variables caused competition to seem like mutualism. In a quasi-experiment that partially controlled competitor abundances with seeding, omitted variables caused competition to seem about 35% weaker than it really was, even though the experiment occurred in an abandoned agricultural field where environmental heterogeneity was expected to be relatively low. Despite decades of research, consistently accurate estimates of competitive interactions remain elusive. The most foolproof way around this problem is true experiments that avoid omitted variable bias by completely controlling competitor abundances, but such experiments are rare.

Soil Aggregate Stability and Grassland Productivity Associations in a Northern Mixed-Grass Prairie.

Soil aggregate stability data are often predicted to be positively associated with measures of plant productivity, rangeland health, and ecosystem functioning. Here we revisit the hypothesis that soil aggregate stability is positively associated with plant productivity. We measured local (plot-to-plot) variation in grassland community composition, plant (aboveground) biomass, root biomass, % water-stable soil aggregates, and topography. After accounting for spatial autocorrelation, we observed a negative association between % water-stable soil aggregates (0.25-1 and 1-2 mm size classes of macroaggregates) and dominant graminoid biomass, and negative associations between the % water-stable aggregates and the root biomass of a dominant sedge (Carex filifolia). However, variation in total root biomass (0-10 or 0-30 cm depths) was either negatively or not appreciably associated with soil aggregate stabilities. Overall, regression slope coefficients were consistently negative thereby indicating the general absence of a positive association between measures of plant productivity and soil aggregate stability for the study area. The predicted positive association between factors was likely confounded by variation in plant species composition. Specifically, sampling spanned a local gradient in plant community composition which was likely driven by niche partitioning along a subtle gradient in elevation. Our results suggest an apparent trade-off between some measures of plant biomass production and soil aggregate stability, both known to affect the land's capacity to resist erosion. These findings further highlight the uncertainty of plant biomass-soil stability associations.

Direct effects dominate responses to climate perturbations in grassland plant communities.

Theory predicts that strong indirect effects of environmental change will impact communities when niche differences between competitors are small and variation in the direct effects experienced by competitors is large, but empirical tests are lacking. Here we estimate negative frequency dependence, a proxy for niche differences, and quantify the direct and indirect effects of climate change on each species. Consistent with theory, in four of five communities indirect effects are strongest for species showing weak negative frequency dependence. Indirect effects are also stronger in communities where there is greater variation in direct effects. Overall responses to climate perturbations are driven primarily by direct effects, suggesting that single species models may be adequate for forecasting the impacts of climate change in these communities.

Fire and grazing effects on wind erosion, soil water content, and soil temperature.

Selective grazing of burned patches can be intense if animal distribution is not controlled and may compound the independent effects of fire and grazing on soil characteristics. Our objectives were to quantify the effects of patch burning and grazing on wind erosion, soil water content, and soil temperature in sand sagebrush (Artemisia filifolia Torr.) mixed prairie. We selected 24, 4-ha plots near Woodward, OK. Four plots were burned during autumn (mid-November) and four during spring (mid-April), and four served as nonburned controls for each of two years. Cattle were given unrestricted access (April-September) to burned patches (<2% of pastures) and utilization was about 78%. Wind erosion, soil water content, and soil temperature were measured monthly. Wind erosion varied by burn, year, and sampling height. Wind erosion was about 2 to 48 times greater on autumn-burned plots than nonburned plots during the dormant period (December-April). Growing-season (April-August) erosion was greatest during spring. Erosion of spring-burned sites was double that of nonburned sites both years. Growing-season erosion from autumn-burned sites was similar to nonburned sites except for one year with a dry April-May. Soil water content was unaffected by patch burn treatments. Soils of burned plots were 1 to 3 degrees C warmer than those of nonburned plots, based on mid-day measurements. Lower water holding and deep percolation capacity of sandy soils probably moderated effects on soil water content and soil temperature. Despite poor growing conditions following fire and heavy selective grazing of burned patches, no blowouts or drifts were observed.