A comparative study of oak (Quercus, Fagaceae) seedling physiology during summer drought in southern California.

Natural recruitment of oaks appears to be declining throughout the northern hemisphere. Summer drought poses a potentially important barrier to oak recruitment in southern California. To evaluate this barrier, we grew evergreen Quercus agrifolia and deciduous Q. lobata from seeds near parental trees. We measured water relations, chlorophyll fluorescence, and gas exchange during these seedlings' fourth and fifth summers and compared them to neighboring adults. Most seedlings had substantially lower values for predawn xylem pressure potential (Ψ(pd)), minimum photosystem II (PSII) quantum efficiency (Φ(PSIIMIN)), maximum quantum efficiency for PSII under dark-adapted leaf conditions (Fv/Fm), and maximum photosynthetic assimilation (Amax), and higher values for maximum nonphotochemical quenching (NPQmax) than did conspecific adults. The high, unvarying Ψ(pd) values of the adults suggest they use perennially available groundwater. Quercus lobata seedlings commonly had lower values for Ψ(pd) than did Q. agrifolia, and values for Ψ(pd) and Φ(PSIIMIN) were significantly related to size in Q. lobata but not in Q. agrifolia. These data suggest important interspecific differences in root architecture. Lower values for Φ(PSIIMIN), Fv/Fm, and higher NPQmax in Q. agrifolia indicate that Q. agrifolia seedlings were usually under more stress than Q. lobata, which typically had higher Amax rates than did Q. agrifolia seedlings. Diurnal photosynthesis curves were quite flat for Q. agrifolia, but they peaked in the morning for Q. lobata. Established seedlings appeared to be under more stress than adults, but this stress did not appear severe enough to cause death. Access to perennially available groundwater may be crucial for the seedling to sapling transition.

Demography and recruitment limitations of three oak species in California.

We review published studies on the demography and recruitment of California oak trees and focus on the widespread dominant species of the foothill woodlands, Quercus douglasii, Q. lobata, and Q. agrifolia, to ascertain the nature and strength of evidence for a decline in populations of these species. The vast majority of studies have been of short duration (less than three years), focused on the acorn and seedling life stages, and conducted at few locations within each species geographic range. We summarize the extensive body of research that has been conducted on the biological and physical factors that limit natural seedling recruitment of oaks. The oak "regeneration problem" has largely been inferred from current stand structure rather than by demographic analyses, which in part reflects the short-term nature of most oak research. When viewed over longer periods of time usingfield surveys or historical photos, the evidence for a regeneration problem in foothill oaks is mixed. Q. douglasii shows very limited seedling or sapling recruitment at present, but longer term studies do not suggest a decline in tree density, presumably because rare recruitment is sufficient to offset low rates of mortality of overstory individuals. Q. agrifolia appears to be stable or increasing in some areas, but decreasing in areas recently impacted by the disease Phytophthora ramorum. Evidence from the few available studies is more consistent in suggesting long-term declines in foothill populations of Q. lobata. Long-term monitoring, age structure analysis, and population models are needed to resolve the current uncertainty over the sustainability of oak woodlands in California.

The effects of neighbors on the growth and survival of shrub seedlings following fire.

Where plant species vie for limited resources, disturbances might preclude competition by releasing a flush of nutrients, or by reducing biomass and thereby diminishing the consumption of resources. However, if new seedlings colonize in clumps, they may still deplete resources within the local aggregations, which may then reduce their growth and survivorship. We investigated competition among seedlings in a burned area by examining the relationship between the performance of newly established shrub seedlings of Ceanothus impressus and (1) the proximity and (2) the identity of their near neighbors. We also investigated the relationship between neighbor proximity and the availability of water. Both survivorship and growth of C. impressus were positively associated with increasing distance to near neighbors, in a manner consistent with resource competition. The availability of water (as determined by pre-dawn xylem pressure potentials) tended to be greater when neighbors were farther away, providing evidence that water was a resource for which plants were competing. This conclusion is reinforced by the finding that the effects of neighbors were stronger in drought years, suggesting that yearly variation in the availability of an important resource (water) can affect the strength of competitive interactions. This suggests that after disturbances, when some resources are apparently abundant on a large scale, competition may be important in determining the small scale patterns of seedling growth and survival.

Invasion of maritime chaparral by the introduced succulentCarpobrotus edulis : The roles of fire and herbivory.

Invasion by the alien succulent,Carpobrotus edulis, has become a common occurrence after fire in maritime chaparral in coastal California, USA. We studied post-burnCarpobrotus establishment in chaparral that lackedCarpobrotus plants before the fire and compared seedbank and field populations in adjacent burned and unburned stands.Carpobrotus seeds were abundant in deer scat and in the soil before burning. Burning did not enhance germination: many seeds were apparently killed by fire and seed bank cores taken after fire revealed no germinable seeds. Laboratory tests showed that temperatures over 105°C for five minutes killedCarpobrotus seeds. In a field experiment involving use of herbivore exclosures, we found that herbivory was an important source of mortality for seedlings in both burned and unburned chaparral. All seedlings, however, died outside of the burn regardless of the presence of cages. Establishment there is apparently limited by factors affecting plant physiology. In the burned area, seedlings that escaped herbivory grew very rapidly. Overall, it appears that herbivory limited seedling establishment in both burned and unburned sites but that the post-burn soil environment supportedCarpobrotus growth in excess of herbivore use, thus promoting establishment.