San Francisco Estuary and Watershed Science

Restorationists commonly plant overstory and understory species simultaneously at the outset of restoration, but a mature forest canopy may be necessary to facilitate survival of light-intolerant understory species. We conducted two experiments in riparian forest restoration sites along the Sacramento River to determine whether: (1) introducing understory species is more successful at the beginning of restoration or after the canopy has developed; (2) canopy cover directly (via reduced light) or indirectly (by reducing non-native competition) facilitates survival of native understory species; and (3) seeding or planting seedlings of understory species is most effective. Seven native understory species were introduced as both seeds and seedlings into an experiment that manipulated canopy cover (open or canopy) and non-native grass competition (control or grass-specific herbicide). We conducted another experiment using shade cloth to directly test the effect of different light levels on seedling survival and growth of three species. Both experiments showed that four species (Aristolochia californica, Carex barbarae, Clematis ligusticifolia, and Vitis californica) had higher survival under low-light conditions (canopy or shade cloth). In contrast, three species (Artemisia douglasiana, Euthamia occidentalis and Rubus californica) had similar survival across open and canopy conditions. Cover of unplanted understory vegetation (mostly non-native) was much lower under the canopy than in open plots treated with grass-specific herbicide. Establishment from seed was generally low and highly variable. Our results suggest that to restore understory species in riparian forests in north–central California: light-intolerant understory species should be planted after canopy closure; canopy cover is more effective than grass-specific herbicide at reducing non-native understory cover; and planting seedlings is more successful than direct seeding.