Woody plants in grasslands: post-encroachment stand dynamics.

Woody plant abundance is widely recognized to have increased in savannas and grasslands worldwide. The lack of information on the rates, dynamics, and extent of increases in shrub abundance is a major source of uncertainty in assessing how this vegetation change has influenced biogeochemical cycles. Projecting future consequences of woody cover change on ecosystem function will require knowledge of where shrub cover in present-day stands lies relative to the realizable maximum for a given soil type within a bioclimatic region. We used time-series aerial photography (1936, 1966, and 1996) and field studies to quantify cover and biomass of velvet mesquite (Prosopis velutina Woot.) following its proliferation in a semidesert grassland of Arizona. Mapping of individual shrubs indicated an encroachment phase characterized by high rates of bare patch colonization. Upon entering a stabilization phase, shrub cover increases associated with recruitment and canopy expansion were largely offset by contractions in canopy area of other shrub patches. Instances of shrub disappearance coincided with a period of below-average rainfall (1936-1966). Overall, shrub cover (mean +/- SE) on sandy uplands with few and widely scattered shrubs in 1902 was dynamically stable over the 1936-1996 period averaging approximately 35% +/- 5%. Shrub cover on clayey uplands in 1936 was 17% +/- 2% but subsequently increased twofold to levels comparable to those on sandy uplands by 1966 (36% +/- 7%). Cover on both soils then decreased slightly between 1966 and 1996 to 28% +/- 3%. Thus, soil properties influenced the rate at which landscapes reached a dynamic equilibrium, but not the apparent endpoint. Although sandy and clayey landscapes appear to have stabilized at comparable levels of cover, shrub biomass was 1.4 times greater on clayey soils. Declines in shrub cover between 1966 and 1996 were accompanied by a shift to smaller patch sizes on both sandy and clayey landscapes. Dynamics observed during the stabilization phase suggest that density-dependent regulation may be in play. If woody cover has transitioned from directional increases to a dynamic equilibrium, biomass projections will require monitoring and modeling patch dynamics and stand structure rather than simply changes in total cover.