Transpiration dynamics support resource partitioning in African savanna trees and grasses

Citation: Holdo, R. M., & Nippert, J. B. (2015). Transpiration dynamics support resource partitioning in African savanna trees and grasses. Ecology, 96(6), 1466-1472. doi:10.1890/14-1986.1It is still far from clear whether and to what extent trees and grasses partition soil moisture in tropical savannas. A major reason for this is that we don't know how snapshot data on rooting differences translate into ecologically relevant patterns of water use at seasonal scales. We used stable isotopes in soil and stem water to quantify functional rooting profiles in grasses and two tree species in a South African savanna. Concurrently, we measured tree sap-flow velocity, grass canopy temperature (a transpiration correlate), and soil moisture content at multiple depths over the course of a growing season. We used lasso regression to identify the dominant soil moisture layers driving daily variation in tree and grass water-use metrics while controlling for weather variables. We found clear rooting depth differences between grasses (shallow) and trees (deep) from the isotopic data, and these patterns were strongly supported by the water-use data, which showed that grasses and trees predominantly responded to soil moisture availability at 5 and 40 cm depth, respectively. Our results provide a rare example of mechanistic support for the resource partitioning hypothesis in savannas, with important implications for our understanding of tree-grass dynamics under altered precipitation regimes

Savanna Tree Seedlings are Physiologically Tolerant to Nighttime Freeze Events

Citation: O'Keefe, K., Nippert, J. B., & Swemmer, A. M. (2016). Savanna Tree Seedlings are Physiologically Tolerant to Nighttime Freeze Events. Frontiers in Plant Science, 7, 13. doi:10.3389/fpls.2016.00046Freeze events can be important disturbances in savanna ecosystems, yet the interactive effect of freezing with other environmental drivers on plant functioning is unknown. Here, we investigated physiological responses of South African tree seedlings to interactions of water availability and freezing temperatures. We grew widely distributed South African tree species (Colophospermum mopane, Combretum apiculaturn, Acacia nigrescens, and Cassia abbreviate) under well watered and water-limited conditions and exposed individuals to nighttime freeze events. Of the four species studied here, C. mopane was the most tolerant of lower water availability. However, all species were similarly tolerant to nighttime freezing and recovered within one week following the last freezing event. We also show that water limitation somewhat increased freezing tolerance in one of the species (C. mopane). Therefore, water limitation, but not freezing temperatures, may restrict the distribution of these species, although the interactions of these stressors may have species specific impacts on plant physiology. Ultimately, we show that unique physiologies can exist among dominant species within communities and that combined stresses may play a currently unidentified role in driving the function of certain species within southern Africa

A Bayesian model for predicting local El Niño events using tree ring widths and cellulose δ18O

This is the peer reviewed version of the following article: Nippert, Jesse B.; Hooten, Mevin B.; Sandquist, Darren R.; Ward, Joy K. (2010). "A Bayesian model for predicting local El Niño events using tree ring widths and cellulose δ18O." Journal of Geophysical Research: Biogeosciences, 115(G1):G01011. http://www.dx.doi.org/10.1029/2009JG001101., which has been published in final form at http://doi.org/10.1029/2009JG001101. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving.The oxygen stable isotopic composition (δ18O) of cellulose recorded in annual tree rings reflects the climate and precipitation history experienced during tree growth and development. Here, we show proxy evidence of El Niño events over the past 30 years using juniper tree rings from southern California, United States. The relationship between tree ring δ18O in α cellulose and annual ring width was negative during most years, reflecting amount-driven fractionation during precipitation. During El Niño years, the relationship between δ18O and ring width was positive with the largest ring widths correlated to the heaviest δ18O. Warmer sea surface temperatures during vapor formation and the strengthening of vapor transport from the eastern Pacific Ocean inland is the most likely mechanism driving heavier δ18O in precipitation during El Niño years. Based on this varying relationship between tree ring width and climate-dependent δ18O values, we created a model to estimate the probability that a given annual tree ring was formed during an El Niño or non–El Niño year. The methods used in this analysis differ from standard dendrochronological technique because we explicitly account for the varying relationship between climate and tree ring characteristic during an El Niño or non–El Niño year. Moreover, our approach accommodates uncertainty in model parameters and predictions better than traditional classification methods. The application of this model to prehistory tree samples or samples of unknown age may allow for El Niño detection and subsequent determination of changes in El Niño frequency

Lack of eutrophication in a tallgrass prairie ecosystem over 27 years

Many North American grasslands are receiving atmospheric nitrogen (N) deposition at rates above what are considered critical eutrophication thresholds. Yet, potential changes in grassland function due to anthropogenic N deposition are poorly resolved, especially considering that other dynamic factors such as land use and precipitation can also affect N availability. To better understand whether elevated N deposition has altered ecosystem structure or function in North American grasslands, we analyzed a 27-year record of ecophysiological, community, and ecosystem metrics for an annually burned Kansas tallgrass prairie. Over this time, despite increasing rates of N deposition that are within the range of critical loads for grasslands, there was no evidence of eutrophication. Plant N concentrations did not increase, soil moisture did not decline, forb diversity did not decline, and the relative abundance of dominant grasses did not shift toward more eutrophic species. Neither aboveground primary productivity nor N availability to plants increased. The fates of deposited N in grasslands are still uncertain, and could include management losses through burning and grazing. However, evidence from this grassland indicates that eutrophication of North American grassland ecosystems is not an inevitable consequence of current levels of N deposition

Intraspecific Trait Variability in Andropogon gerardii, a Dominant Grass Species in the US Great Plains

The climatic conditions in the North American Great Plains are highly variable, characteristic of an inter-continental climate. Antecedent climate history has impacted the flora of Great Plains grasslands, resulting in high species richness as well as dominance by only a few grass species, such as Andropogon gerardii. While the productivity of A. gerardii is well described, the individual physiological, and morphological characteristics that confer species dominance over wide spatial gradients are not clearly understood. We performed a literature search to assess intra-specific trait variability of A. gerardii from as many locations as possible. Ultimately, only 13 locations in the Great Plains have reported common plant functional traits (PFTs) for this species. To best represent site-specific climate conditions, plant functional trait data (8 PFTs) were collected from literature reporting ambient growing conditions, and excluded experimental manipulations. For most PFTs, we found insufficient data to fully quantify the range of variation across the geographical extent of A. gerardii dominance. This is surprising given that we focused on the most abundant grass in one of the most well-studied regions globally. Furthermore, trait data collected from our literature search showed a high degree of variability, but no strong relationships were observed between mean trait values and climate predictors. Our review of the literature on A. gerardii suggests a role for trait variability as a mechanism enabling the dominance of this species across large regions such as the Great Plains of North America

The interactive effects of press/pulse intensity and duration on regime shifts at multiple scales

Citation: Ratajczak, Z., D'Odorico, P., Collins, S. L., Bestelmeyer, B. T., Isbell, F. I., & Nippert, J. B. (2017). The interactive effects of press/pulse intensity and duration on regime shifts at multiple scales. Ecological Monographs, 87(2), 198-218. doi:10.1002/ecm.1249Regime shifts are difficult-to-reverse transitions that occur when an ecosystem reorganizes around a new set of self-reinforcing feedbacks. Regime shifts are predicted to occur when the intensity of some exogenous driver variable, such as temperature, annual harvest rate, or nutrient addition rate, gradually approaches and crosses a threshold value, initiating a transition to an alternative state. However, many driver variables now change rapidly as presses or pulses, not gradually, requiring new conceptual frameworks for understanding and predicting regime shifts. We argue that identifying and controlling regime shifts in response to presses and pulses will require a greater focus on the duration, not just the intensity, of changes in driver variables. In ecosystems with slower dynamics, transitions to an alternative state can take years to decades and as a result, a driver press with an intensity capable of resulting in a regime shift over long time spans may fail to cause a regime shift when applied for shorter durations. We illustrate these ideas using simulations of local-scale alternative stable state models and preliminary evidence from long-term grazing and eutrophication experiments. The simulations also suggest that small changes in the duration of driver presses or pulses can determine whether an ecosystem recovers to its original state. These insights may extend to larger scales. In spatially extended simulations that included patchiness, spatial heterogeneity, and spatial connectivity, all patches recovered to their original state after shorter presses. However, once press duration exceeded a threshold, growing proportions of the landscape shifted to an alternative state as press duration increased. We observed similar patchy transitions in a catchment-scale experiment that reinstated frequent fires approximately halfway through a regime shift from grassland to shrubland, initiated by fire suppression. In both the local-and larger-scale models, the threshold duration needed to elicit regime shifts decreased as press intensity increased or when factors counteracting regime shifts weakened. These multiple lines of evidence suggest that conceptualizing regime shifts as an interactive function of the intensity and duration of driver changes will increase understanding of the varying effects of driver presses, pulses, and cycles on ecosystem dynamics

Assessing the Roles of Fire Frequency and Precipitation in Determining Woody Plant Expansion in Central U.S. Grasslands

Woody plant expansion into grasslands and savannas is occurring and accelerating worldwide and often impacts ecosystem processes. Understanding and predicting the environmental and ecological impacts of encroachment has led to a variety of methodologies for assessing its onset, transition, and stability, generally relying on dynamical systems approaches. Here we continue this general line of investigation to facilitate the understanding of the roles of precipitation frequency and intensity and fire frequency on the conversion of grasslands to woody‐dominated systems focusing on the central United States. A low‐dimensional model with stochastic precipitation and fire disturbance is introduced to examine the complex interactions between precipitation and fire as mechanisms that may suppress or facilitate increases in woody cover. By using Lyapunov exponents, we are able to ascertain the relative control exerted on woody encroachment through these mechanisms. Our results indicate that precipitation frequency is a more important control on woody encroachment than the intensity of individual precipitation events. Fire, however, exerts a much more dominant impact on the limitation of encroachment over the range of precipitation variability considered here. These results indicate that fire management may be an effective strategy to slow the onset of woody species into grasslands. While climate change might predict a reduced potential for woody encroachment in the near future, these results indicate a reduction in woody fraction may be unlikely when considering anthropogenic fire suppression

Response of a mixed grass prairie to an extreme precipitation event

Citation: Concilio, A. L., Prevey, J. S., Omasta, P., O'Connor, J., Nippert, J. B., & Seastedt, T. R. (2015). Response of a mixed grass prairie to an extreme precipitation event. Ecosphere, 6(10), 12. doi:10.1890/es15-00073.1Although much research has been conducted to measure vegetation response to directional shifts in climate change drivers, we know less about how plant communities will respond to extreme events. Here, we evaluate the response of a grassland community to an unprecedented 43 cm rainfall event that occurred in the Front Range of Colorado in September, 2013 using vegetation plots that had been monitored for response to simulated precipitation changes since 2011. This rain caused soils to stay at or above field capacity for multiple days, and much of the seed bank germinated following the early autumn event. Annual introduced grasses, especially cheatgrass (Bromus tectorum), and several introduced forbs demonstrated strong positive increases in cover the following growing season. Native cool season grasses and native forbs showed limited changes in absolute cover despite continued high soil water availability, while native warm season grasses increased in cover the following summer. Treatments that previously altered the amounts and seasonality of rainfall during the 2011-2013 interval showed legacy effects impacting cover responses of introduced species and warm-season native grasses. Resin bag estimates of inorganic nitrogen flux resulting from the event indicated twice as much nitrogen movement compared to any previous collections during the 2011-2013 interval. Nitrogen additions to a subset of plots made in spring of 2014 demonstrated that the relative cover of introduced species could be further increased with additional soil nitrogen. Collectively, these results support the contention that extreme precipitation events can favor species already benefiting from other environmental change drivers

Evidence of physiological decoupling from grassland ecosystem drivers by an encroaching woody shrub

Shrub encroachment of grasslands is a transformative ecological process by which native woody species increase in cover and frequency and replace the herbaceous community. Mechanisms of encroachment are typically assessed using temporal data or experimental manipulations, with few large spatial assessments of shrub physiology. In a mesic grassland in North America, we measured inter- and intra-annual variability in leaf δ¹³C in Cornus drummondii across a grassland landscape with varying fire frequency, presence of large grazers and topographic variability. This assessment of changes in individual shrub physiology is the largest spatial and temporal assessment recorded to date. Despite a doubling of annual rainfall (in 2008 versus 2011), leaf δ¹³C was statistically similar among and within years from 2008-11 (range of −28 to −27‰). A topography*grazing interaction was present, with higher leaf δ¹³C in locations that typically have more bare soil and higher sensible heat in the growing season (upland topographic positions and grazed grasslands). Leaf δ¹³C from slopes varied among grazing contrasts, with upland and slope leaf δ¹³C more similar in ungrazed locations, while slopes and lowlands were more similar in grazed locations. In 2011, canopy greenness (normalized difference vegetation index – NDVI) was assessed at the centroid of individual shrubs using high-resolution hyperspectral imagery. Canopy greenness was highest mid-summer, likely reflecting temporal periods when C assimilation rates were highest. Similar to patterns seen in leaf δ¹³C, NDVI was highest in locations that typically experience lowest sensible heat (lowlands and ungrazed). The ability of Cornus drummondii to decouple leaf physiological responses from climate variability and fire frequency is a likely contributor to the increase in cover and frequency of this shrub species in mesic grassland and may be generalizable to other grasslands undergoing woody encroachment

Fire dynamics distinguish grasslands, shrublands and woodlands as alternative attractors in the Central Great Plains of North America

This review synthesizes evidence that altered fire frequency drives discontinuous ecosystem transitions from mesic grasslands to shrublands or woodlands in the Central Great Plains, USA. Long-term fire manipulations reveal that grassland to shrubland transitions are triggered when fire-free intervals increase from 1–3 years to ≥ 3–8 years, and longer fire returns (~10 years or more) result in transitions to woodlands. Grazing and soil properties alter these fire thresholds. Grassland to shrubland transitions are abrupt and exhibit nonlinear relationships between driver and state variables. Transitions to shrublands and woodlands exhibit hysteresis, where reintroducing frequent fires does not reverse transitions in management-relevant time-scales (decades). Nonlinear transitions and hysteresis emerge because grasses generate positive feedbacks with fire that create strong demographic barriers for shrub and tree establishment. Fire-free intervals allow shrubs and trees to reach a size sufficient to survive fire, reproduce and disrupt the fire feedback loop through competition. Synthesis: Mesic grasslands, shrublands and woodlands constitute self-reinforcing states (alternative attractors) separated by critical fire frequency thresholds. Even without major shifts in climate, altered fire frequency can produce dramatic state changes, highlighting the importance of fire for predicting future ecosystem states. Local management should focus on prevention of unwanted transitions rather than post hoc restoration