Summary1. Rangeland ecologists have been debating the validity of two current paradigms for the evaluation of vegetation dynamics on rangelands. This debate frequently contrasts the conventional model of continuous and reversible vegetation dynamics (range model) with a more contemporary model that can accommodate discontinuous and non-reversible vegetation change (state-and-transition model). 2. The range and the state-and-transition models are conceptually related to the equilibrium and non-equilibrium paradigms within ecology, respectively. The methodological dichotomy that has developed between the range and the state-and-transition models has fostered the perception that these two ecological paradigms are mutually exclusive. We challenge this perception and contend that both methodologies and their corresponding paradigms are non-exclusive. 3. Equilibrium and non-equilibrium ecosystems are not distinguished on the basis of unique processes or functions, but rather by the evaluation of system dynamics at various temporal and spatial scales. Consequently, ecosystems may express both equilibrium and non-equilibrium dynamics. This confirms early interpretations that ecosystems are distributed along a continuum from equilibrium to non-equilibrium states. 4. Although both equilibrium and non-equilibrium dynamics occur in numerous ecosystems, the empirical evidence is frequently confounded by (i) uncertainty regarding the appropriate evidence necessary to distinguish between paradigms; (ii) disproportionate responses among vegetation attributes to climate and grazing; (iii) comparisons among systems with varying degrees of managerial involvement; and (iv) the evaluation of vegetation dynamics at various spatial and temporal scales. 5. Synthesis and applications . This critique supports the conclusion that a paradigm shift has not taken place in rangeland ecology, but rather, the debate has forced a more comprehensive interpretation of vegetation dynamics along the entirety of the equilibrium-non-equilibrium continuum. Therefore, the rangeland debate should be redirected from the dichotomy between paradigms to one of paradigm integration.
This article synthesizes the ecological concepts and perspectives underpinning the development and application of stateand-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. The Texas and Oklahoma Agricultural Experiment Stations and USDA-NRI Ecosystem Program (92-37101-7463) funded field research that contributed to this synthesis and the stations provided partial support to the authors during manuscript preparation.
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. Resumen La meta de esta síntesis es iniciar el desarrollo de un marco teó rico unificado para la evaluació n de los umbrales que sea capaz de relacionar la teoría ecoló gica y procesos con el conocimiento de manejo y su aplicació n. Los objetivos específicos incluyen la investigació n de los mecanismos de los umbrales, la elaboració n de los componentes de los umbrales, la introducció n de categorías y trayectorias de los umbrales y la presentació n de una definició n operativa de los umbrales ecoló gicos. Un mayor entendimiento de los umbrales ecoló gicos es esencial porque ellos han venido a ser el punto focal dentro del marco de la teoría de estados estables y transició n, y su ocurrencia tiene consecuencias criticas para el manejo de los pastizales. La ocurrencia de umbrales puede ser mejor interpretada como un cambio de la dominancia de reacciones negativas que mantienen la resilencia del ecosistema a la dominancia de reacciones positivas que degradan la resilencia y promueven el desarrollo de estados poster...
A 44-yr record of herbaceous vegetation change was analysed for three contrasting grazing regimes within a semi-arid savanna to evaluate the relative contribution of confined livestock grazing and climatic variability as agents of vegetation change. Grazing intensity had a significant, directional effect on the relative composition of short-and midgrass response groups; their composition was significantly correlated with time since the grazing regimes were established. Interannual precipitation was not significantly correlated with response group composition. However, interannual precipitation was significantly correlated with total plant basal area while time since imposition of grazing regimes was not, but both interannual precipitation and time since the grazing regimes were established were significantly correlated with total plant density. Vegetation change was reversible even though the herbaceous community had been maintained in an altered state for ca. 60 yr by intensive livestock grazing. However, ca. 25 yr were required for the mid-grass response group to recover following the elimination of grazing and recovery occurred intermittently. The increase in mid-grass composition was associated with a significant decrease in total plant density and an increase in mean individual plant basal area. Therefore, we failed to reject the hypotheses based on the proportional change in relative response group composition with grazing intensity and the distinct effects of grazing and climatic variability on response group composition, total basal area and plant density. Long-term vegetation change indicates that grazing intensity established the long-term directional change in response group composition, but that episodic climate events defined the short-term rate and trajectory of this change and determines the upper limit on total basal area. The occurrence of both directional and non-directional vegetation responses were largely a function of (1) the unique responses of the various community attributes monitored and (2) the distinct temporal responses of these community attributes to grazing and climatic variation. This interpretation supports previous conclusions that individual ecosystems may exist in equilibrial and non-equilibrial states at various temporal and spatial scales.
basal area dynamics of dominant graminoid species were analyzed across three grazing intensity treatments (heavily grazed, moderately grazed and ungrazed) at the Texas A&M University Agricultural Research Station on the Edwards Plateau, Texas. Grazing intensity was identified as the primary influence on long-term variations in species composition. Periodic weather events, including a severe drought (1951)(1952)(1953)(1954)(1955)(1956), had little direct influence on composition dynamics. However, the drought interacted with grazing intensity in the heavily grazed treatment to exacerbate directional changes caused by grazing intensity.Species response to grazing was individualistic and noisy. Three response groups were identified. Taller, more productive mid-grasses were most abundant under moderate or no grazing. Short grasses were most abundant under heavy grazing. Intermediate species were most abundant under moderate grazing and opportunistic to weather patterns. Graminoid diversity increased with the removal or reduction of grazing intensity. The moderately and ungrazed treatments appeared most resistant to short-term weather fluctuations, while the heavily grazed treatment demonstrated significant resilience when grazing intensity was reduced after over 110 yr of overgrazing.Identification of a 'climax' state is difficult. Significant directional change, which took nearly 20 yr, appears to continue in the ungrazed treatment after 45 yr of succession. The observed, relatively linear patterns of perennial grass composition within the herbaceous patches of this savanna were generally explained by traditional Clementsian succession. However, when dynamics of the herbaceous community are combined with the woody component of this savanna, the frequency and intensity of fire becomes more important. Across the landscape, successional changes follow several pathways. When vegetation change is influenced by several factors, a multi-scale model is necessary to demonstrate interactions and feedbacks and accurately describe successional patterns. Absence of fires, with or without grazing, leads ultimately to a Juniperus/Quercus woodland with grazing intensity primarily influencing the fuel load and hence fire intensity.
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