Patterns of understory diversity in mixed coniferous forests of southern California impacted by air pollution

The forests of the San Bernardino Mountains have been subject to ozone and nitrogen (N) deposition for some 60 years. Much work has been done to assess the impacts of these pollutants on trees, but little is known about how the diverse understory flora has fared. Understory vegetation has declined in diversity in response to elevated N in the eastern U.S. and Europe. Six sites along an ozone and N deposition gradient that had been part of a long-term study on response of plants to air pollution beginning in 1973 were resampled in 2003. Historic ozone data and leaf injury scores confirmed the gradient. Present-day ozone levels were almost half of these, and recent atmospheric N pollution concentrations confirmed the continued air pollution gradient. Both total and extractable soil N were higher in sites on the western end of the gradient closer to the urban source of pollution, pH was lower, and soil carbon (C) and litter were higher. The gradient also had decreasing precipitation and increasing elevation from west to east. However, the dominant tree species were the same across the gradient. Tree basal area increased during the 30-year interval in five of the sites. The two westernmost sites had 30-45% cover divided equally between native and exotic understory herbaceous species, while the other sites had only 3-13% cover dominated by native species. The high production is likely related to higher precipitation at the western sites as well as elevated N. The species richness was in the range of 24 to 30 in four of the sites, but one site of intermediate N deposition had 42 species, while the easternmost, least polluted site had 57 species. These were primarily native species, as no site had more than one to three exotic species. In three of six sites, 20-40% of species were lost between 1973 and 2003, including the two westernmost sites. Two sites with intermediate pollution had little change in total species number over 30 years, and the easternmost site had more species in 2003. The easternmost site is also the driest and has the most sunlight filtering to the forest floor, possibly accounting for the higher species richness. The confounding effects of the precipitation gradient and possibly local disturbances do not show a simple correlation of air pollution with patterns of native and invasive species cover and richness. Nevertheless, the decline of native species and dominance by exotic species in the two westernmost polluted sites is cause for concern that air pollution is affecting the understory vegetation adversely.