Invasion by Non-Native Annual Grasses: The Importance of Species Biomass, Composition, and Time Among California Native Grasses of the Central Valley

The Central Valley of California is noted for its dearth of remnant native grass populations and for low native grass seedling establishment within grasslands now dominated by non‐native annual species. In contrast, remnant populations are common along the coast, and studies have shown an ability for seedlings and adults to compete with non‐native annual grasses. The invasibility of well‐established populations of native grasses in the Central Valley remains unclear. The objectives of this study were to compare the invasibility of native grasses differing in density and species composition and, given the species in this study, to assess the ability of mixes with greater species richness to resist invasion relative to their abilities in monoculture. In the Sacramento Valley of California, six species of native grasses were planted at three densities in monospecific and mixed‐species plots. Percent cover of native perennial and non‐native annual grasses was measured in years 2 and 3, and biomass was sampled in year 5. Native grass biomass and, to a lesser extent, species composition were important in explaining variation in non‐native grass invasibility in the fifth year. Species‐rich treatments did not experience less invasion than would be expected by the proportional invasibility of each species in monoculture. However, invasibility of plots consisting of slower growing, shorter statured species decreased over time, suggesting a successional benefit to diverse communities. This study demonstrates that established stands of native grasses in the Sacramento Valley can resist invasion by non‐native annual grasses and that stand biomass is a particularly important factor in determining invasibility.     

Effects of Established Perennial Grasses on Introduction of Native Forbs in California

Prairie restoration is not complete without the establishment of both grasses and forbs. However, if desirable forbs and grasses are seeded simultaneously, control of broadleaf weeds is problematic. If possible, a two‐step process of introducing forbs after establishing grasses would allow use of broadleaf‐specific herbicides at the critical early stages of grass growth. We conducted experiments to investigate methods for introducing forbs into previously restored native perennial grasslands on rural roadsides in the Sacramento Valley of California. In one experiment, we studied the effects of background vegetation (established perennial grasses or tilled ground) on seven native forb species planted from seed. In a second experiment, we evaluated the effects of background vegetation (existing perennial grasses or tilled ground) and container size (36 ml or 105 ml) with excavation technique (excavation by core removal [core] or by creating an impression [dibble]) on the growth of transplants of the native perennial forbs Asclepias fascicularis and Sisyrinchium bellum. The presence of established perennial grasses reduced the growth of seeded forbs, but did not affect transplants, indicating the vulnerability of seedling forbs to interference. When compared to control plots that had been tilled in the autumn, weed canopy cover was significantly lower in the presence of perennial grasses if seeded with forbs, but not in the presence of perennial grasses alone. Both transplanted species grew better in the large container/core treatment than the small container/dibble treatment; however, existing grasses eliminated these positive effects. Asclepias fascicularis performed better when grown in large containers than in small containers, but its growth was not affected by excavation method; S. bellum performed better when planted with the core method than the dibble method of excavation, but container size made no difference. We attribute differences in the responses of the species to interactions between phenological differences and expansive clay soils that naturally de‐compact upon drying.     

Restoration of California Native Grasses and Clovers: The Roles of Clipping,Broadleaf Herbicide,and Native Grass Density

One of the major challenges confronting grassland restoration of highly invaded communities is increasing the diversity of native species. There is surprisingly little research investigating how reconstructed native grasslands respond to common management techniques and how these techniques influence the relative establishment of both native grasses and forbs. Despite the diversity and wide distribution of native clovers in California, few practitioners incorporate them into grassland restoration plans. Conversely, non‐native clovers have been seeded extensively onto California rangelands. This study addresses the following questions: (1) Using readily available management tools, is there a strategy that can benefit the growth of both planted native bunchgrasses and seeded clovers? (2) Do native bunchgrasses compete with establishing clovers and non‐native grasses? (3) Do native and non‐native clovers differ in their response to management treatments or in their productivity? Plots were established to test three factors in different combinations over 3 years: (1) early spring clipping, (2) initial broadleaf herbicide, and (3) native bunchgrass planting density. Native and non‐native clovers were seeded in years 2 and 3. Early spring clipping did not have a significant effect on native bunchgrass cover, yet it did result in greater growth of native and non‐native clovers. The direction of the response to broadleaf herbicide changed between years for native bunchgrasses and was consistently negative for native clovers. Plots with higher native grass densities did not adversely affect the seeded clovers, yet non‐native grass cover was reduced. Native and non‐native clovers exhibited similar responses to clipping and established at similar densities.     

Effects of Competition and Life History Stage on the Expression of Local Adaptation in Two Native Bunchgrasses

Concerns about the use of genetically appropriate material in restoration often focus on questions of local adaptation. Many reciprocal transplant studies have demonstrated local adaptation in native plant species, but very few have examined how interspecific competition affects the expression of adaptive variation. Our study examined regional scales of adaptation between foothill and coastal populations of two California native bunchgrasses (Elymus glaucus and Nassella pulchra). By combining competitive manipulations with reciprocal transplants, we examined the importance of the vegetation at a site as a selective factor in the process of local adaptation. By monitoring survival and reproduction of reciprocally transplanted populations over the course of 3 years, we also studied the effect of life history stage on the expression of local adaptation. For most of the fitness components we measured, local adaptation was detected and interspecific competition consistently amplified its expression. Expression of local adaptation was especially apparent in the more inbreeding species E. glaucus and suggests that with weaker gene flow, selection may be more effective in creating ecotypes within this species. Local adaptation was detected at all life history stages but was most strongly expressed in traits associated with adult reproduction and the viability of seeds produced by the transplants. Taken together, our results indicate that the importance of local adaptation will become more apparent in the later stages of a restoration project as the plants at a site begin to reproduce and as they experience greater interspecific competition from the maturing vegetation at the site.     

Reducing Competitive Suppression of a Rare Annual Forb by Restoring Native California Perennial Grasslands

Populations of the rare annual forb Amsinckia grandiflora may be declining because of competitive suppression by exotic annual grasses, and may perform better in a matrix of native perennial bunchgrasses. We conducted a field competition experiment in which Amsinckia seedlings were transplanted into forty 0.64‐m² experimental plots of exotic annual grassland or restored perennial grassland. The perennial grassland plots were restored using mature 3 cm‐diameter plants of the native perennial bunchgrass Poa secunda planted in three densities. The exotic annual grassland plots were established in four densities through manual removal of existing plants. Both grass types reduced soil water potential with increasing biomass, but this reduction was not significantly different between grass types. Both grass types significantly reduced the production of Amsinckia inflorescences. At low and intermediate densities (dry biomass per unit area of 20–80 g/m²), the exotic annual grasses reduced Amsinckia inflorescence number to a greater extent than did Poa, although at high densities (>90 g/m²) both grass types reduced the number of Amsinckia inflorescences to the same extent. The response of Amsinckia inflorescence number to Poa biomass was linear, whereas the same response to the annual grass biomass is logarithmic, and appeared to be related to graminoid cover. This may be because of the different growth forms exhibited by the two grass types. Results of this research suggest that restored native perennial grasslands at intermediate densities have a high habitat value for the potential establishment of the native annual A. grandiflora.