David D. Briske

Consequences of More Extreme Precipitation Regimes for Terrestrial Ecosystems

ABSTRACT Amplification of the hydrological cycle as a consequence of global warming is forecast to lead to more extreme intra-annual precipitation regimes characterized by larger rainfall events and longer intervals between events. We present a conceptual framework, based on past investigations and ecological theory, for predicting the consequences of this underappreciated aspect of climate change. We consider a broad range of terrestrial ecosystems that vary in their overall water balance. More extreme rainfall regimes are expected to increase the duration and severity of soil water stress in mesic ecosystems as intervals between rainfall events increase. In contrast, xeric ecosystems may exhibit the opposite response to extreme events. Larger but less frequent rainfall events may result in proportional reductions in evaporative losses in xeric systems, and thus may lead to greater soil water availability. Hydric (wetland) ecosystems are predicted to experience reduced periods of anoxia in response to prolonged intervals between rainfall events. Understanding these contingent effects of ecosystem water balance is necessary for predicting how more extreme precipitation regimes will modify ecosystem processes and alter interactions with related global change drivers.

Rotational Grazing on Rangelands: Reconciliation of Perception and Experimental Evidence

Abstract In spite of overwhelming experimental evidence to the contrary, rotational grazing continues to be promoted and implemented as the only viable grazing strategy. The goals of this synthesis are to 1) reevaluate the complexity, underlying assumptions, and ecological processes of grazed ecosystems, 2) summarize plant and animal production responses to rotational and continuous grazing, 3) characterize the prevailing perceptions influencing the assessment of rotational and continuous grazing, and 4) attempt to direct the profession toward a reconciliation of perceptions advocating support for rotational grazing systems with that of the experimental evidence. The ecological relationships of grazing systems have been reasonably well resolved, at the scales investigated, and a continuation of costly grazing experiments adhering to conventional research protocols will yield little additional information. Plant production was equal or greater in continuous compared to rotational grazing in 87% (20 of 23) of the experiments. Similarly, animal production per head and per area were equal or greater in continuous compared to rotational grazing in 92% (35 of 38) and 84% (27 of 32) of the experiments, respectively. These experimental data demonstrate that a set of potentially effective grazing strategies exist, none of which have unique properties that set one apart from the other in terms of ecological effectiveness. The performance of rangeland grazing strategies are similarly constrained by several ecological variables establishing that differences among them are dependent on the effectiveness of management models, rather than the occurrence of unique ecological phenomena. Continued advocacy for rotational grazing as a superior strategy of grazing on rangelands is founded on perception and anecdotal interpretations, rather than an objective assessment of the vast experimental evidence. We recommend that these evidence-based conclusions be explicitly incorporated into management and policy decisions addressing this predominant land use on rangelands.

State-and-Transition Models, Thresholds, and Rangeland Health: A Synthesis of Ecological Concepts and Perspectives

Abstract This article synthesizes the ecological concepts and perspectives underpinning the development and application of state-and-transition models, thresholds, and rangeland health. Introduction of the multiple stable state concept paved the way for the development of these alternative evaluation procedures by hypothesizing that multiple stable plant communities can potentially occupy individual ecological sites. Vegetation evaluation procedures must be able to assess continuous and reversible as well as discontinuous and nonreversible vegetation dynamics because both patterns occur and neither pattern alone provides a complete assessment of vegetation dynamics on all rangelands. Continuous and reversible vegetation dynamics prevail within stable vegetation states, whereas discontinuous and nonreversible dynamics occur when thresholds are surpassed and one stable state replaces another. State-and-transition models can accommodate both categories of vegetation dynamics because they represent vegetation change along several axes, including fire regimes, weather variability, and management prescriptions, in addition to the succession-grazing axis associated with the traditional range model. Ecological thresholds have become a focal point of state-and-transition models because threshold identification is necessary for recognition of the various stable plant communities than can potentially occupy an ecological site. Thresholds are difficult to define and quantify because they represent a complex series of interacting components, rather than discrete boundaries in time and space. Threshold components can be categorized broadly as structural and functional based on compositional and spatial vegetation attributes, and on modification of ecosystem processes, respectively. State-and-transition models and rangeland health procedures have developed in parallel, rather than as components of an integrated framework, because the two procedures primarily rely on structural and functional thresholds, respectively. It may be prudent for rangeland professionals to consider the introduction of these alternative evaluation procedures as the beginning of a long-term developmental process, rather than as an end point marked by the adoption of an alternative set of standardized evaluation procedures.

A Unified Framework for Assessment and Application of Ecological Thresholds

Abstract The goal of this synthesis is to initiate development of a unified framework for threshold assessment that is able to link ecological theory and processes with management knowledge and application. Specific objectives include the investigation of threshold mechanisms, elaboration of threshold components, introduction of threshold categories and trajectories, and presentation of an operational definition of ecological thresholds. A greater understanding of ecological thresholds is essential because they have become a focal point within the state-and-transition framework and their occurrence has critical consequences for land management. Threshold occurrence may be best interpreted as a switch from the dominance of negative feedbacks that maintain ecosystem resilience to the dominance of positive feedbacks that degrade resilience and promote the development of post-threshold states on individual ecological sites. Threshold categories have been identified to serve as ecological benchmarks to describe the extent of threshold progression and increase insight into feedback mechanisms that determine threshold reversibility. Threshold trajectories describe the developmental pathway that post-threshold states may follow once a threshold has been exceeded. These trajectories may produce a continuum of potential post-threshold states, but the majority of them may be organized into four broad states. This framework lends itself to management application by providing an operational definition of thresholds that is based on a probabilistic interpretation. Probabilities associated with 1) the occurrence of triggers that initiate threshold progression, 2) the trajectory of post-threshold states, and 3) threshold reversibility will provide an operational procedure for threshold assessment and application. If thresholds are to play a central role in rangeland ecology and management, then the rangeland profession must accept responsibility for their conceptual development, ecological validity, and managerial effectiveness.

RECOMMENDATIONS FOR DEVELOPMENT OF RESILIENCE-BASED STATE-AND-TRANSITION MODELS

Abstract The objective of this paper is to recommend conceptual modifications for incorporation in state-and-transition models (STMs) to link this framework explicitly to the concept of ecological resilience. Ecological resilience describes the amount of change or disruption that is required to transform a system from being maintained by one set of mutually reinforcing processes and structures to a different set of processes and structures (e.g., an alternative stable state). In light of this concept, effective ecosystem management must focus on the adoption of management practices and policies that maintain or enhance ecological resilience to prevent stable states from exceeding thresholds. Resilience management does not exclusively focus on identifying thresholds per se, but rather on within-state dynamics that influence state vulnerability or proximity to thresholds. Resilience-based ecosystem management provides greater opportunities to incorporate adaptive management than does threshold-based management because thresholds emphasize limits of state resilience, rather than conditions that determine the probability that these limits will be surpassed. In an effort to further promote resilience-based management, we recommend that the STM framework explicitly describe triggers, at-risk communities, feedback mechanisms, and restoration pathways and develop process-specific indicators that enable managers to identify at-risk plant communities and potential restoration pathways. Two STMs representing different ecological conditions and geographic locations are presented to illustrate the incorporation and application of these recommendations. We anticipate that these recommendations will enable STMs to capture additional ecological information and contribute to improved ecosystem management by focusing attention on the maintenance of state resilience in addition to the anticipation of thresholds. Adoption of these recommendations may promote valuable dialogue between researchers and ecosystem managers regarding the general nature of ecosystem dynamics.

Herbivore‐Induced Species Replacement in Grasslands: Is it Driven by Herbivory Tolerance or Avoidance?

Herbivore-induced shifts in species composition have been documented from grasslands throughout the world, but the mechanism(s) of species replacement remains largely unexplored. An experiment was conducted in a transplant garden, on the campus of Texas AM however, tiller number per plant was suppressed by B. saccharoides, but not by S. leucotricha or conspecific neighbors. Defoliation of S. scoparium plants, but not neighbors, negatively impacted the late-seral plants. Selective defoliation of S. sco- parium plants significantly reduced tiller variables of mean mass, leaf blade area, and leaf number, but did not significantly reduce plant variables including mean basal area, tiller number, or annual shoot production. Defoliation of both S. scoparium plants and neighbors increased annual shoot production, mean basal area per plant, mean tiller leaf area, leaf number, tiller mass, stomatal conductance to H20 vapor, and plant xylem pressure potential in comparison with S. scoparium plants grown with comparable, nondefoliated neighbors. An increase in both plant and tiller variables in defoliated S. scoparium plants grown with uniformly defoliated neighbors establishes that replace- ment of a late-seral dominant is not driven by a greater relative expression of herbivory tolerance of mid-seral species. These results collectively suggest that the late-seral dominant, S. scoparium, pos- sesses a greater competitive ability and a comparable or greater degree of herbivory tolerance than the mid-seral species that comprise the community. Therefore, the initial hypothesis was rejected. It can be inferred that the alternative mechanism, selective herbivory of the late-seral dominant, is the dominant mechanism contributing to species replacement. Herbivore-induced modifications of competitive interactions are most like- ly to drive species replacement in grasslands characterized by high and consistent resource availability. This may partially explain why condition and trend analysis was developed and initially implemented in the true and mixed prairie associations of North America and why it is widely used by rangeland managers in these grasslands.

Does grazing mediate soil carbon and nitrogen accumulation beneath C4, perennial grasses along an environmental gradient?

An experiment was conducted to evaluate the influence of long-term (>25 yrs) grazing on soil organic carbon (SOC) and total soil nitrogen (N) accumulation beneath individual plants of three perennial grasses along an environmental gradient in the North American Great Plains. The zone of maximum SOC and N accumulation was restricted vertically to the upper soil depth (0-5 cm) and horizontally within the basal area occupied by individual caespitose grasses, which contributed to fine-scale patterning of soil heterogeneity. Long-term grazing mediated SOC and N accumulation in the tall-, mid- and shortgrass communities, but the responses were community specific. SOC and N were lower beneath Schizachyrium scoparium plants in long-term grazed sites of the tall- and midgrass communities, but higher beneath Bouteloua gracilis plants in the long-term grazed site of the shortgrass community. SOC, but not N, was greater in soils beneath compared to between S. scoparium plants in an abandoned field seeded in 1941, indicating that this caespitose grass accumulated SOC more rapidly than N. SOC and N were greater in the 0-5 cm soil depth beneath a caespitose grass (S. scoparium) compared to a rhizomatous grass (Panicum virgatum) in the tallgrass community, with no significant accumulation of either SOC or N beneath P. virgatum plants. Grazing appears to indirectly mediate nutrient accumulation beneath caespitose grasses along the environmental gradient by modifying the size class distribution of plants. Populations with a greater proportion of large plants have a greater potential for biomass incorporation into soils and may more effectively capture redistributed organic matter from between plant locations. Contrasting plant responses to grazing at various locations along the environmental gradient conform to the predictions of the generalized grazing model, as the selection pressures of grazing and aridity may have also influenced the ability of caespitose grasses to accumulate nutrients in soils beneath them by mediating grazing resistance, competitive ability and population structure.

Isoprene emission from terrestrial ecosystems in response to global change: minding the gap between models and observations

Coupled surface–atmosphere models are being used with increased frequency to make predictions of tropospheric chemistry on a ‘future’ earth characterized by a warmer climate and elevated atmospheric CO2 concentration. One of the key inputs to these models is the emission of isoprene from forest ecosystems. Most models in current use rely on a scheme by which global change is coupled to changes in terrestrial net primary productivity (NPP) which, in turn, is coupled to changes in the magnitude of isoprene emissions. In this study, we conducted measurements of isoprene emissions at three prominent global change experiments in the United States. Our results showed that growth in an atmosphere of elevated CO2 inhibited the emission of isoprene at levels that completely compensate for possible increases in emission due to increases in aboveground NPP. Exposure to a prolonged drought caused leaves to increase their isoprene emissions despite reductions in photosynthesis, and presumably NPP. Thus, the current generation of models intended to predict the response of isoprene emission to future global change probably contain large errors. A framework is offered as a foundation for constructing new isoprene emission models based on the responses of leaf biochemistry to future climate change and elevated atmospheric CO2 concentrations.

Origin, Persistence, and Resolution of the Rotational Grazing Debate: Integrating Human Dimensions Into Rangeland Research

Abstract The debate regarding the benefits of rotational grazing has eluded resolution within the US rangeland profession for more than 60 yr. This forum examines the origin of the debate and the major reasons for its persistence in an attempt to identify common ground for resolution, and to search for meaningful lessons from this central chapter in the history of the US rangeland profession. Rotational grazing was a component of the institutional and scientific response to severe rangeland degradation at the turn of the 20th century, and it has since become the professional norm for grazing management. Managers have found that rotational grazing systems can work for diverse management purposes, but scientific experiments have demonstrated that they do not necessarily work for specific ecological purposes. These interpretations appear contradictory, but we contend that they can be reconciled by evaluation within the context of complex adaptive systems in which human variables such as goal setting, experiential knowledge, and decision making are given equal importance to biophysical variables. The scientific evidence refuting the ecological benefits of rotational grazing is robust, but also narrowly focused, because it derives from experiments that intentionally excluded these human variables. Consequently, the profession has attempted to answer a broad, complex question—whether or not managers should adopt rotational grazing—with necessarily narrow experimental research focused exclusively on ecological processes. The rotational grazing debate persists because the rangeland profession has not yet developed a management and research framework capable of incorporating both the social and biophysical components of complex adaptive systems. We recommend moving beyond the debate over whether or not rotational grazing works by focusing on adaptive management and the integration of experiential and experimental, as well as social and biophysical, knowledge to provide a more comprehensive framework for the management of rangeland systems.

Climate Change and North American Rangelands: Trends, Projections, and Implications

Abstract The amplified “greenhouse effect” associated with increasing concentrations of greenhouse gases has increased atmospheric temperature by 1°C since industrialization (around 1750), and it is anticipated to cause an additional 2°C increase by mid-century. Increased biospheric warming is also projected to modify the amount and distribution of annual precipitation and increase the occurrence of both drought and heat waves. The ecological consequences of climate change will vary substantially among ecoregions because of regional differences in antecedent environmental conditions; the rate and magnitude of change in the primary climate change drivers, including elevated carbon dioxide (CO2), warming and precipitation modification; and nonadditive effects among climate drivers. Elevated atmospheric CO2 will directly stimulate plant growth and reduce negative effects of drying in a warmer climate by increasing plant water use efficiency; however, the CO2 effect is mediated by environmental conditions, especially soil water availability. Warming and drying are anticipated to reduce soil water availability, net primary productivity, and other ecosystem processes in the southern Great Plains, the Southwest, and northern Mexico, but warmer and generally wetter conditions will likely enhance these processes in the northern Plains and southern Canada. The Northwest will warm considerably, but annual precipitation is projected to change little despite a large decrease in summer precipitation. Reduced winter snowpack and earlier snowmelt will affect hydrology and riparian systems in the Northwest. Specific consequences of climate change will be numerous and varied and include modifications to forage quantity and quality and livestock production systems, soil C content, fire regimes, livestock metabolism, and plant community composition and species distributions, including range contraction and expansion of invasive species. Recent trends and model projections indicate continued directional change and increasing variability in climate that will substantially affect the provision of ecosystem services on North American rangelands.