Dynamics of shrub encroachment in an African savanna: relative influences of fire, herbivory, rainfall and density dependence.
Shrub encroachment has been widely observed in savanna regions. This study analysed the causes of shrub encroachment in the lowveld savanna of north-eastern Swaziland, and highlighted management regimes that can be used to reduce or prevent it. The rates and dynamics of shrub encroachment were quantified for the period 1947-90 using aerial photographs, and for 1997 using a ground survey. Five similar areas with different land use histories were compared to investigate the relative importance of fire, herbivory, rainfall, soil type and shrub density in driving shrub dynamics. In the study area as a whole, shrub cover increased from a mean of 2% in 1947 to 31% in 1990. Dichrostachys cinerea accounted for most of the increase in cover, contributing 81% to total shrub cover during 1997. Shrub cover was strongly correlated with shrub density and weakly negatively correlated with tree cover. Shrub encroachment varied across land use fence lines. The key determinants of shrub dynamics were grazing, through its negative effect on fire frequency and an interaction between drought frequency and high shrub cover. Browsing pressure had a significant but minor impact on dynamics, while soil type had no significant effect. High grazing pressures through their effect on fire frequency were critical throughout the study period in promoting shrub encroachment, while the interaction between drought and high shrub cover produced declines in the later stages. Browsing had an impact on encroachment only in the early stages. Frequent fires, facilitated by low grazing pressures, were capable of preventing shrub encroachment. When coupled with drought, frequent fires could reduce high shrub densities. As cover and density were strongly correlated, it can be inferred from the negative correlation between change in cover (density) and initial cover (density) that the rate of shrub encroachment was cover (density)-dependent and that there was a shrub equilibrium of 40% cover, approximating to 2400 plants ha-1. Shrub population growth was driven by events (fire, drought) as well as continuous agents (density dependence, mean browsing and grazing pressure). Bush encroachment can be reversed by a combination of management (frequent fires) and climatic events (drought). The implications for savanna management are discussed.