Long‐term changes in semi‐arid vegetation: Invasion of an exotic perennial grass has larger effects than rainfall variability

Abstract. Questions: This paper examines the long‐term change in the herbaceous layer of semi‐arid vegetation since grazing ceased. We asked whether (1) there were differences in the temporal trends of abundance among growth forms of plants; (2) season of rainfall affected the growth form response; (3) the presence of an invasive species influenced the abundance and species richness of native plants relative to non‐invaded plots, and (4) abundance of native plants and/or species richness was related to the time it took for an invasive species to invade a plot. Location: Alice Springs, Central Australia. Methods: Long‐term changes in the semi‐arid vegetation of Central Australia were measured over 28 years (1976–2004) to partition the effects of rainfall and an invasive perennial grass. The relative abundance (biomass) of all species was assessed 25 times in each of 24 plots (8 m × 1 m) across two sites that traversed floodplains and adjacent foot slopes. Photo‐points, starting in 1972, were also used to provide a broader overview of a landscape that had been intensively grazed by cattle and rabbits prior to the 1970s. Species’abundance data were amalgamated into growth forms to examine their relationship with environmental variation in space and time. Environmental variables included season and amount of rainfall, fire history, soil variability and the colonization of the plots by the exotic perennial grass Cenchrus ciliaris (Buffel grass). Results: Constrained ordination showed that season of rainfall and landscape variables relating to soil depth strongly influenced vegetation composition when Cenchrus was used as a covariate. When Cenchrus was included in constrained ordination, it was strongly related to the decline of all native growth forms over time. Univariate comparisons of non‐invaded vs impacted plots over time revealed unequivocal evidence that Cenchrus had caused the decline of all native growth form groups and species richness. They also revealed a contrasting response of native plants to season of rainfall, with a strong response of native grasses to summer rainfall and forbs to winter rainfall. In the presence of Cenchrus these responses were strongly attenuated. Discussion: Pronounced changes in the composition of vegetation were interpreted as a response to removal of grazing pressure, fluctuations in rainfall and, most importantly, invasion of an exotic grass. Declines in herbaceous species abundance and richness in the presence of Cenchrus appear to be directly related to competition for resources. Indirect effects may also be causing the declines of some woody species from changed fire regimes as a result of increased fuel loads. We predict that Cenchrus will begin to alter landscape level processes as a result of the direct and indirect effects of Cenchrus on the demography of native plants when there is a switch from resource limited (rainfall) establishment of native plants to seed limited recruitment.     

The importance of environment vs. disturbance in the vegetation mosaic of Central Arizona

Abstract. The vegetation of central Arizona is a mosaic of four vegetation types: chaparral, chaparral grassland, woodland, and woodland grassland. We analysed ten environmental variables, three disturbance variables, and five disturbance indicators to answer the question: What is the relative importance of environment and disturbance in explaining the vegetation pattern of our study area? We found that chaparral, chaparral grassland, and woodland are differentiated primarily by environmental factors and have high stability in the landscape. In contrast, woodland grassland is differentiated primarily by disturbance and is likely an early‐successional stage of woodlands. Although other researchers have indicated that semi‐arid vegetation is generally unstable, the vegetation of central Arizona is composed of two systems: those with a more stable landscape position determined primarily by environmental factors and those with a less stable landscape position determined primarily by disturbance factors.     

Plant colonization windows in a mesic old field succession

Abstract. Closed canopy vegetation often prevents the colonization of plant species. Therefore the majority of plant species are expected to appear at the initial phase of post‐agricultural succession in mesic forest environment with moderate levels of resources. This hypothesis was tested with data from the Buell‐Small Successional Study, NJ, USA, one of the longest continuous fine‐scale studies of old‐field succession. The study started in 1958, including old fields with different agricultural histories, landscape contexts, and times of abandonment. In each year of the study, the cover values of plant species were recorded in 48 permanent plots of 1 m² in each field. We analysed the temporal patterns of colonization at plot scale and related these to precipitation data and other community characteristics. The number of colonizing species decreased significantly after ca. 5 yr, coinciding with the development of a continuous canopy of perennial species. However, species turnover remained high throughout the whole successional sequence. The most remarkable phenomenon is the high inter‐annual variation of all studied characteristics. We found considerable temporal collapses of vegetation cover that were synchronized among fields despite their different developmental stages and distinctive species compositions. Declines of total cover were correlated with drought events. These events were associated with peaks of local species extinctions and were followed by increased colonization rates. The transitions of major successional stages were often connected to these events. We suggest that plant colonization windows opened by extreme weather events during succession offer optimum periods for intervention in restoration practice.     

Herbaceous vegetation change in variable rangeland environments: The relative contribution of grazing and climatic variability

Abstract. 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 mid‐grass 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.