Invasive insect effects on nitrogen cycling and host physiology are not tightly linked.

Invasive insects may dramatically alter resource cycling and productivity in forest ecosystems. Yet, although responses of individual trees should both reflect and affect ecosystem-scale responses, relationships between physiological- and ecosystem-scale responses to invasive insects have not been extensively studied. To address this issue, we examined changes in soil nitrogen (N) cycling, N uptake and allocation, and needle biochemistry and physiology in eastern hemlock (Tsuga canadensis (L) Carr) saplings, associated with infestation by the hemlock woolly adelgid (HWA) (Adelges tsugae Annand), an invasive insect causing widespread decline of eastern hemlock in the eastern USA. Compared with uninfested saplings, infested saplings had soils that exhibited faster nitrification rates, and more needle (15)N uptake, N and total protein concentrations. However, these variables did not clearly covary. Further, within infested saplings, needle N concentration did not vary with HWA density. Light-saturated net photosynthetic rates (Asat) declined by 42% as HWA density increased from 0 to 3 adelgids per needle, but did not vary with needle N concentration. Rather, Asat varied with stomatal conductance, which was highest at the lowest HWA density and accounted for 79% of the variation in Asat. Photosynthetic light response did not differ among HWA densities. Our results suggest that the effects of HWA infestation on soil N pools and fluxes, (15)N uptake, needle N and protein concentrations, and needle physiology may not be tightly coupled under at least some conditions. This pattern may reflect direct effects of the HWA on N uptake by host trees, as well as effects of other scale-dependent factors, such as tree hydrology, affected by HWA activity.

Oak seedling growth and ectomycorrhizal colonization are less in eastern hemlock stands infested with hemlock woolly adelgid than in adjacent oak stands.

Invasive, non-indigenous, phytophagous insects have caused widespread declines in several dominant tree species. The decline in dominant tree species may lead to cascading effects on other tree and microbial species and their interactions, affecting forest recovery following the decline. In the eastern USA, eastern hemlock (Tsuga canadensis (L.) Carr) is declining because of infestation by the hemlock woolly adelgid (HWA; Adelges tsugae Annand). Northern red oak (Quercus rubra L.) is a common replacement species in declining hemlock stands, but reduced mycorrhizal inoculum potential in infested hemlock stands may cause oak to grow more slowly compared with oak in oak stands. We grew red oak seedlings for one growing season in declining hemlock-dominated stands infested with HWA and in adjacent oak-dominated stands. Ectomycorrhizal root tip density and morphotype richness in soil cores were 63 and 27% less, respectively, in declining hemlock stands than in oak stands. Similarly, ectomycorrhizal percent colonization and morphotype richness on oak seedlings were 33 and 30% less, respectively, in declining hemlock stands than in oak stands. In addition, oak seedlings in declining hemlock stands had 29% less dry mass than oak seedlings in oak stands. Analysis of covariance indicated that morphotype richness could account for differences in oak seedling dry mass between declining hemlock stands and oak stands. Additionally, oak seedling dry mass in declining hemlock stands significantly decreased with decreasing ectomycorrhizal percent colonization and morphotype richness. These results suggest that oak seedling growth in declining hemlock stands is affected by reduced ectomycorrhizal inoculum potential. Further, the rate of forest recovery following hemlock decline associated with HWA infestation may be slowed by indirect effects of HWA on the growth of replacement species, through effects on ectomycorrhizal colonization and morphotype richness.

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.

Tracing atmospheric nitrate deposition in a complex semiarid ecosystem using delta17O.

The isotopic composition of nitrate collected from aerosols, fog, and precipitation was measured and found to have a large 17O anomaly with delta17O values ranging from 20 percent per thousand to 30% percent per thousand (delta17O = delta17O - 0.52(delta18O)). This 17O anomaly was used to trace atmospheric deposition of nitrate to a semiarid ecosystem in southern California. We demonstrate that the delta17O signal is a conserved tracer of atmospheric nitrate deposition and is a more robust indicator of N deposition relative to standard delta18O techniques. The data indicate that a substantial portion of nitrate found in the local soil, stream, and groundwater is of atmospheric origin and does not undergo biologic processing before being exported from the system.