Spartina alterniflora was first introduced into south San Francisco Bay in the 1970’s. Since that time it has spread to new areas within the south bay and is especially well established at four sites. The spread of this introduced species was evaluated by comparing its vegetative and reproductive characteristics to the native cordgrass, Spartina foliosa. The characters studied were intertidal distribution, phenology, aboveground and belowground biomass, growth rates, seed production, and germination rates. Spartina alterniflora has a wider intertidal distribution than S. foliosa and outproduced the native cordgrass in all aspects that were studied. These results indicate that the introduced species has a much better chance of becoming established in new areas than the native species, and once established, it spreads more rapidly vegetatively than the native species. Spartina alterniflora is likely to continue to spread to new areas in the bay and displace the native plant. In addition, this introduced species may effect sedimentation dynamics, available detritus, benthic algal production, wrack deposition and disturbance, habitat structure for native wetland animals, benthic invertebrate populations, and shorebird and wading bird foraging areas. *** DIRECT SUPPORT *** A01BY058 00013

The introduction and spread of smooth cordgrass ( Spartina alterniflora ) in South San Francisco Bay

Analysis of spatial pattern of the seed bank of an old field in Maryland showed that the seed bank was composed primarily of annual and short—lived perennial species. Tree and shrub species were poorly represented. Seed abundance decreased with depth in the soil. The seeds of all species generally had clustered distributions in the soil. However, there was no trend in mean cluster size within or among species, or at different soil depths. The lack of definite trends indicates that the spatial distributions of seeds are governed primarily by environmental factors, and to a lesser degree by biological factors. The method of spatial pattern analysis developed for this study represents a significant improvement over traditional contiguous quadrat methods. In this method, individuals' positions are mapped in the field using cartesian coordinates, and the quadrat data are subsequently derived mathematically. The resolution of pattern is greatly improved, and several problems that reduce the effectiveness of other methods are eliminated. This method can be used to advantage in any study in which spatial patterns analysis is performed, particularly if the objects can be mapped easily. A comparison of three sampling techniques revealed that the precision of seed—number estimates can be improved by taking a large number of small samples, or by subsampling large whole units with very small subunits. The smaller number of large samples favored by many seed—bank researchers generally results in imprecise seed estimates. This result has implications for spatial studies of any clustered organism.

Spatial Pattern Analysis of Seed Banks: An Improved Method and Optimized Sampling

Spartina alterniflora was first introduced into south San Francisco Bay in the 1970's. Since that time it has spread to new areas within the south bay and is especially well established at four sites. The spread of this introduced species was evaluated by comparing its vegetative and reproductive characteristics to the native cordgrass, Spartina foliosa. The characters studied were intertidal distribution, phenology, aboveground and belowground biomass, growth rates, seed production, and germination rates. Spartina alterniflora has a wider intertidal distri- bution than S. foliosa and outproduced the native cordgrass in all aspects that were studied. These results indicate that the introduced species has a much better chance of becoming established in new areas than the native species, and once established, it spreads more rapidly vegetatively than the native species. Spartina alterniflora is likely to continue to spread to new areas in the bay and displace the native plant. In addition, this introduced species may affect sedimentation dynamics, available detritus, benthic algal production, wrack deposition and disturbance, habitat structure for native wetland animals, benthic invertebrate populations, and shorebird and wading bird foraging areas.

The Introduction and Spread of Smooth Cordgrass (Spartina alterniflora) in

Although tidal wetland vegetation patterns are typically related to elevation, we hypothesized that the vertical range of a species may shift where the topography is more heterogeneous. We examined plant species occurrences in relation to elevation, proximity to the bay, and proximity to tidal creeks at a near-pristine wetland in San Quintin Bay, Baja California, Mexico. At the whole-wetland scale, most species occurred primarily within a 30-cm elevation band (the marsh plain). However, Spartina foliosa occurred only at the bayward margin, even though “suitable” elevations were present further inland. A similar pattern was found in San Diego Bay. At the microtopographic scale, three species on the marsh plain were strongly influenced by elevation, whereas four species responded to both elevation and proximity to tidal creeks. The latter species tended to “avoid” the lower 10 cm of the marsh plain except near a tidal creek. Species richness was thus greater (by 0.6 species at the lowest 10-cm class) at the tidal creek margin. Better drainage near creeks is the hypothesized cause. Our results help explain why species that are transplanted to constructed wetlands do not always grow at the full range of elevations they occupy in natural wetlands. We recommend that species be introduced to their modal elevation (determined from nearby reference marshes) and that salt-marsh construction designs include topographic heterogeneity (complex tidal creek networks). The analysis of broad-scale and fine-scale patterns of occurrence also suggests new habitat nomenclature. Elevation-based terms (“low,”“middle,” and “high” marsh) should be replaced by a system that considers elevation, landscape position, and conspicuous species. We suggest three habitat designations: (a) the high marsh—a 30- to 70-cm elevation range with Salicornia subterminalis; (b) the marsh plain—a 30-cm elevation range with heterogeneous topography and up to nine common species; and (c) cordgrass habitat—the bayward portion of the marsh plain and lower elevations, all occupied by Spartina foliosa. Although these habitats do not have discrete boundaries, separate terms are needed for wetland restoration plans and these designations will improve recognition that vegetation patterns respond to horizontal, as well as vertical, position.

Californian Salt-Marsh Vegetation: An Improved Model of Spatial Pattern

Seedling recruitment in salt marsh plant communities is generally precluded in dense vegetation by competition from adults, but is also relatively rare in disturbance-generated bare space. We examined the constraints on seedling recruitment in New England salt marsh bare patches. Under typical bare patch conditions seed germination is severely limited by high substrate salinities. We examined the germination requirements of common high marsh plants and found that except for one notably patch-dependent fugitive species, the germination of high marsh plants is strongly inhibited by the high soil salinities routinely encountered in natural bare patches. Watering high marsh soil in the greenhouse to alleviate salt stress resulted in the emergence of up to 600 seedlings/225 cm2. The vast majority of this seed bank consisted of Juncus gerardi, the only common high marsh plant with high seed set. We tested the hypothesis that salt stress limits seedling contributions to marsh patch secondary succession in the field. Watering bare patches with fresh water partially alleviated patch soil salinities and dramatically increased both the emergence and survival of seedlings. Our results show that seedling recruitment by high marsh perennial turfs is limited by high soil salinities and that consequently their population dynamics are determined primarily by clonal growth processes. In contrast, populations of patch-dependent fugitive marsh plants which cannot colonize vegetatively are likely governed by spatially and temporally unpredictable windows of low salinities in bare patches.

Salt stress limitation of seedling recruitment in a salt marsh plant community.

Abondance, composition specifique et distribution des graines enfouies dans les marais sales de la Baie de San Francisco

Dale R. Hopkins

Papers

A study of the seed bank of a salt marsh in Northern San Francisco Bay