Causes and consequences of woody plant encroachment into western North American grasslands

Abstract

As woody plants encroach into grasslands, grass biomass, density and cover decline as wood plant biomass, density and cover increase. There is also a shift in location of the biomass from mostly belowground in the grasslands to aboveground in the woodlands. In addition, species richness and diversity change as herbaceous species are replaced by woody species. This is not a new phenomenon, but has been going on continually as the climate of the Planet has changed. However, in the past 160 years the changes have been unparalleled. The process is encroachment not invasion because woody species that have been increasing in density are native species and have been present in these communities for thousands of years. These indigenous or native woody species have increased in density, cover and biomass because of changes in one or more abiotic or biotic factors or conditions. Woody species that have increased in density and cover are not the cause of the encroachment, but the result of changes of other factors. Globally, the orbit of the Earth is becoming more circular and less elliptical, causing moderation of the climate. Additional global climate changing factors including elevated levels of CO 2 and parallel increases in temperature are background factors and probably not the principal causes directing the current wave of encroachment. There is probably not a single reason for encroachment, but a combination of factors that are difficult to disentangle. The prime cause of the current and recent encroachment appears to be high and constant levels of grass herbivory by domestic animals. This herbivory reduces fine fuel with a concomitant reduction in fire frequency or in some cases a complete elimination of fire from these communities. Conditions would now favor the woody plants over the grasses. Reduced grass competition, woody plant seed dispersal and changes in animal populations seem to modify the rate of encroachment rather than being the cause. High concentrations of atmospheric CO 2 are not required to explain current woody plant encroachment. Changes in these grassland communities will continue into the future but the specifics are difficult to predict. Density, cover and species composition will fluctuate and will probably continue to change. Increased levels of anthropogenic soil nitrogen suggest replacement of many legumes by other woody species. Modification and perhaps reversal of the changes in these former grassland communities will be an arduous, continuing and perhaps impossible management task.