| Gradual 
              replacement of meadow by forest species.  Among 
              meadow species cover declined steeply with establishment of grand 
              fir and associated reductions in light (figure, below: upper left 
              panel), but richness (upper right panel) declined more slowly. 
            Forest 
            herbs colonized rapidly (within decades), and within 60-80 yr dominated 
            the understory (lower panels). Richness 
              and cover of forest understory species showed weaker relationships 
              with overstory structure than did richness and cover of meadow species 
              (regression model R2 for forest species: 0.34-0.42; 
              model R2 for meadow species: 0.48-0.54). Conclusions 
               
                | Rapid 
                  replacement of meadow species |   
                | _15%20Jul%202007_lg.jpg) |   
                | 
                    
                      | Forest 
                        understory plants rapidly colonize beneath young stands 
                        of grand fir. |  |  With conifer 
              establishment, replacement of meadow by forest species is rapid. 
              The pace of conversion, reinforced by positive feedbacks between 
              trees and soils, may be indicative of a shift to an alternative 
              stable state. Loss 
              of cover and richness of meadow species (and limited presence of 
              these species in the seed 
              bank) may pose barriers to restoration of native meadows.  Removing 
              trees during the earliest stages of encroachment may be the most 
              effective strategy for maintaining these ecosystems. |