Bush Lupine Mortality, Altered Resource Availability, and Alternative Vegetaion States

Nitrogen-fixing plants, by altering the availability of soil N, potentially facilitate plant invasion. Here we describe how herbivore-driven mortality of a native Nfixing shrub, bush lupine (Lupinus arboreus), increases soil N and light availability, which promotes invasion by introduced grasses to the detriment of a native plant community. Soils under live and dead lupine stands contained large amounts of total N, averaging 3.14 mg N/g dry mass of soil (398 g/m2) and 3.45 mg N/g dry mass of soil (438 g/m2), respectively, over four years. In contrast, similar lupine-free soil was low in N and averaged only 1.66 mg N/g dry mass of soil (211 g/m2) over three years. The addition of N fertilizer to lupine-free soil produced an 81% increase in aboveground plant biomass compared to plots unamended with N. Mean rates of net N mineralization were higher under live lupine and where mass die-off of lupine had occurred compared to soils free of bush lupine. At all sites, only 2.5–4.2% of the total soil N pool was mineralized annually. Soil enriched by lupine is not available to colonists while lupines are alive. The dense canopy of lupine shades soil under shrubs, reducing average photon-flux density in late spring from 1725 mmol·m22·s21 (full sunlight) to 13 mmol·m22·s21 (underneath shrubs). Stand die-off due to insect herbivory exposed this bare, enriched soil. In January, when annual plants are establishing, average photon-flux density under dead lupines killed by insect herbivores was 370 mmol·m22·s21, compared to the photon-flux density under live lupines of the same age, which averaged 83 mmol·m22·s21. The availability of bare, N-rich patches of soil enabled nonnative annuals (primarily Lolium multiflorum and Bromus diandrus) to colonize sites, grow rapidly, and dominate the plant assemblage until lupines reestablished after several years. The N content of these grasses was significantly greater than the N content of the mostly native plants that occupied adjacent coastal prairie devoid of bush lupine. Between 57 and 70% of the net amount of N mineralized annually was taken up by introduced grasses and subsequently returned to the soil upon the death of these annuals. Even in the absence of further N inputs, we estimate that it would take at least 25 yr to reduce the soil N pool by 50%, indicating that the reestablishment of the native prairie flora is likely to be long term.