John C. Callaway

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

ABSTRACT 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 Quintín 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.

Declining Biodiversity: Why Species Matter and How Their Functions Might Be Restored in Californian Tidal Marshes

S diversity is being lost in habitats that are increasingly diminished by development, fragmentation, and urban runoff; the sensitive species drop out and a few aggressive ones persist, at the expense of others. Alarmed by declining biodiversity, many conservationists and researchers are asking what happens to ecosystem functioning if we lose species, how diverse communities can be restored, which (if any) particular species are critical for performing ecosystem services, and which functions are most critical to ecosystem sustainability. In southern California, 90% of the coastal wetland area has been destroyed, and remaining wetlands continue to be damaged; even the region’s protected reserves are threatened by highway and utility-expansion projects. The fate of biodiversity in these diminished wetlands serves to warn other regions of the need for continual assessment of the status and function of both common and rare species, as well as the need for experimental tests of their importance—before they are lost. This article synthesizes data for tidal marshes of the Californian biogeographic region, which stretches from Point Conception near Santa Barbara south to Bahia San Quintin in Baja California. We focus on the broad marsh plain, which is dominated by eight species of halophytes (salt-tolerant plants; Figure 1). From regional censuses, we document the recent loss of short-lived species from several wetlands. From eyewitness accounts of tidal-exclusion events at Estero de Punta Banda and Tijuana Estuary, we link species loss to the interruption of tidal influence. And from experimental plantings of marsh halophytes in a bare restoration site, we document the difficulty of restoring plant diversity, demonstrate Joy B. Zedler, Aldo Leopold Chair of Restoration Ecology, Botany Department and Arboretum, 430 Lincoln Drive, University of Wisconsin– Madison, Madison, WI 53706, is a wetland ecologist who has studied southern California coastal wetlands for about 30 years. John C. Callaway, assistant professor in the Department of Environmental Science, University of San Francisco, San Francisco, CA 94117, conducts field research at Tijuana Estuary and San Francisco Bay; he is a wetland ecologist whose research focuses on sediment and vegetation dynamics in restored wetlands. Gary Sullivan is a wetland ecologist with experience in freshwater lakes and streams, estuaries, and salt marshes; his current focus is on restoring large wetlands along the Illinois River for a nonprofit organization, The Wetlands Initiative, Chicago, IL 60604-3703.

SPECIES-RICH PLANTINGS INCREASE BIOMASS AND NITROGEN ACCUMULATION IN A WETLAND RESTORATION EXPERIMENT

Our test of the hypothesis that biomass and nitrogen would increase with more species-rich plantings simultaneously vegetated a salt marsh restoration site and dem- onstrated that on average, randomly chosen, 6-species plantings accumulated more biomass and nitrogen than the mean for 0- and 1-species assemblages, with the mean for 3-species assemblages being intermediate. In addition, we found that individual species (from the pool of eight native halophytes) differed in their functional capacity, with Salicornia vir- ginica (Sv) and Jaumea carnosa contributing the greatest biomass when planted alone, while Triglochin concinna had the highest tissue N concentrations. When planted alone, Sv accumulated comparable amounts of biomass and nitrogen as in the multispecies plots, indicating that individual species can have a large effect on particular functions. Soil TKN in the surface 0-5 cm was greater in 6-species plots than unplanted plots in 1999, while both 3- and 6-species plots were greater than unplanted plots in 2000; however, there were no differences at 5-20 cm depth and no species-specific effects. Root and shoot biomass both increased with species richness, with total biomass of 6-species plots averaging 995.6 ? 120.5 g/m2 in 2000, compared to the mean for 1-species plots (572.1 + 90.3 g/m2) and unplanted plots (164.5 ? 24.7 g/m2). Still, at the age of three years, root biomass was only about one-third that of the species-rich reference site, and shoot biomass was one-half to one-fifth the maxima reported for reference salt marshes. Species-specific effects were found for Sv, which had high biomass of both roots and shoots in the multispecies plots (55% of aboveground biomass in 3-species plots and 41% in 6-species plots) and the highest pool of N (52% of the N pool in 3-species plots and 42% in 6-species plots), even though only one-eighth of the initial plantings were Sv. However, when plots with this species were excluded from the analysis, the species-richness effect persisted. Thus, ecosystem function, as measured by biomass and N accumulation, increased with species richness regardless of dominance by the highly productive Sv. We conclude that manipulating the richness and composition of plantings offers ecosystem restorationists an effective tool for accelerating the rate of functional development.

Evaluating Tidal Marsh Sustainability in the Face of Sea-Level Rise: A Hybrid Modeling Approach Applied to San Francisco Bay

Background Tidal marshes will be threatened by increasing rates of sea-level rise (SLR) over the next century. Managers seek guidance on whether existing and restored marshes will be resilient under a range of potential future conditions, and on prioritizing marsh restoration and conservation activities. Methodology Building upon established models, we developed a hybrid approach that involves a mechanistic treatment of marsh accretion dynamics and incorporates spatial variation at a scale relevant for conservation and restoration decision-making. We applied this model to San Francisco Bay, using best-available elevation data and estimates of sediment supply and organic matter accumulation developed for 15 Bay subregions. Accretion models were run over 100 years for 70 combinations of starting elevation, mineral sediment, organic matter, and SLR assumptions. Results were applied spatially to evaluate eight Bay-wide climate change scenarios. Principal Findings Model results indicated that under a high rate of SLR (1.65 m/century), short-term restoration of diked subtidal baylands to mid marsh elevations (−0.2 m MHHW) could be achieved over the next century with sediment concentrations greater than 200 mg/L. However, suspended sediment concentrations greater than 300 mg/L would be required for 100-year mid marsh sustainability (i.e., no elevation loss). Organic matter accumulation had minimal impacts on this threshold. Bay-wide projections of marsh habitat area varied substantially, depending primarily on SLR and sediment assumptions. Across all scenarios, however, the model projected a shift in the mix of intertidal habitats, with a loss of high marsh and gains in low marsh and mudflats. Conclusions/Significance Results suggest a bleak prognosis for long-term natural tidal marsh sustainability under a high-SLR scenario. To minimize marsh loss, we recommend conserving adjacent uplands for marsh migration, redistributing dredged sediment to raise elevations, and concentrating restoration efforts in sediment-rich areas. To assist land managers, we developed a web-based decision support tool (www.prbo.org/sfbayslr).

Modeling Tidal Marsh Distribution with Sea-Level Rise: Evaluating the Role of Vegetation, Sediment, and Upland Habitat in Marsh Resiliency

Tidal marshes maintain elevation relative to sea level through accumulation of mineral and organic matter, yet this dynamic accumulation feedback mechanism has not been modeled widely in the context of accelerated sea-level rise. Uncertainties exist about tidal marsh resiliency to accelerated sea-level rise, reduced sediment supply, reduced plant productivity under increased inundation, and limited upland habitat for marsh migration. We examined marsh resiliency under these uncertainties using the Marsh Equilibrium Model, a mechanistic, elevation-based soil cohort model, using a rich data set of plant productivity and physical properties from sites across the estuarine salinity gradient. Four tidal marshes were chosen along this gradient: two islands and two with adjacent uplands. Varying century sea-level rise (52, 100, 165, 180 cm) and suspended sediment concentrations (100%, 50%, and 25% of current concentrations), we simulated marsh accretion across vegetated elevations for 100 years, applying the results to high spatial resolution digital elevation models to quantify potential changes in marsh distributions. At low rates of sea-level rise and mid-high sediment concentrations, all marshes maintained vegetated elevations indicative of mid/high marsh habitat. With century sea-level rise at 100 and 165 cm, marshes shifted to low marsh elevations; mid/high marsh elevations were found only in former uplands. At the highest century sea-level rise and lowest sediment concentrations, the island marshes became dominated by mudflat elevations. Under the same sediment concentrations, low salinity brackish marshes containing highly productive vegetation had slower elevation loss compared to more saline sites with lower productivity. A similar trend was documented when comparing against a marsh accretion model that did not model vegetation feedbacks. Elevation predictions using the Marsh Equilibrium Model highlight the importance of including vegetation responses to sea-level rise. These results also emphasize the importance of adjacent uplands for long-term marsh survival and incorporating such areas in conservation planning efforts.

Evolution of tidal creek networks in a high sedimentation environment: A 5-year experiment at Tijuana Estuary, California

In a large (8 ha) salt marsh restoration site, we tested the effects of excavating tidal creeks patterned after reference systems. Our purposes were to enhance understanding of tidal creek networks and to test the need to excavate creeks during salt marsh restoration. We compared geomorphic changes in areas with and without creek networks (n = 3; each area 1.3 ha) and monitored creek cross-sectional areas, creek lengths, vertical accretion, and marsh surface elevations for 5 yr that included multiple sedimentation events. We hypothesized that cells with creeks would develop different marsh surface and creek network characteristics (i.e., surface elevation change, sedimentation rate, creek cross-sectional area, length, and drainage density). Marsh surface vertical accretion averaged 1.3 cm yr−1 with large storm inputs, providing the opportunity to assess the response of the drainage network to extreme sedimentation rates. The constructed creeks initially filled due to high accretion rates but stabilized at cross-sectional areas matching, or on a trajectory toward, equilibrium values predicted by regional regression equations. Sedimentation on the marsh surface was greatest in low elevation areas and was not directly influenced by creeks. Time required for cross-sectional area stabilization ranged from 0 to > 5 yr, depending on creek order. First-order constructed creeks lengthened rapidly (mean rate of 1.3 m yr−1) in areas of low elevation and low vegetation cover. New (volunteer) creeks formed rapidly in cells without creeks in areas with low elevation, low vegetation cover, and high elevation gradient (mean rate of 6.2 m yr−1). After 5 yr, volunteer creeks were, at most, one-fourth the area of constructed creeks and had not yet reached the upper marsh plain. In just 4 yr, the site’s drainage density expanded from 0.018 to reference levels of 0.022 m m−2. Pools also formed on the marsh plain due to sediment resuspension associated with wind-driven waves. We conclude that excavated creeks jump-started the development of drainage density and creek and channel dimensions, and that the tidal prism became similar to those of the reference site in 4–5 yr.

Relationship between topographic heterogeneity and vegetation patterns in a Californian salt marsh

Abstract Questions: Are species richness and species abundances higher in the presence of tidal creeks? Do species richness and species abundances vary with plot size? Location: Intertidal plain of Volcano Marsh, Bahia de San Quintin, Mexico. Methods: We analysed vegetation patterns in large areas (cells) with tidal creeks (+creek) and without (−creek). We surveyed vegetation cover, microtopography, habitat type, and distance to creeks in nested plots of five sizes, 0.1, 0.25, 1, 2.5, and 10 m2. Results: Species richness, frequency, cover, and assemblages differed between ±creek cells. Richness tended to be higher in +creek cells, and cover and frequency of individual species differed significantly between ±creek cells. We found consistent patterns in vegetation structure across plot sizes. We encountered 13 species that occurred in 188 unique assemblages. The most common assemblage had six species: Batis maritima, Frankenia salina, Salicornia bigelovii, S. virginica, Salicornia spec. and Triglochin concinna. This assemblage occurred in ±creek cells and at all spatial scales. Of the most common assemblages all but one were composed of multiple species (3–9 species/plot). Conclusions: The persistence of vegetation patterns across a 100-fold range in spatial scale suggests that similar environmental factors operate broadly to determine species establishment and persistence. Differences in assemblage composition result from variation of frequency and cover of marsh plain species, particularly Suaeda esteroa and Monanthochloe littoralis. The recommendation for restoration of Californian salt marshes is to target (and plant) multi-species assemblages, not monocultures. Nomenclature: Hickman (1993).

Diversity–function relationships changed in a long-term restoration experiment

The central tenet of biodiversity-ecosystem function (BEF) theory, that species richness increases function, could motivate restoration practitioners to incorporate a greater number of species into their projects. But it is not yet clear how well BEF theory predicts outcomes of restoration, because it has been developed through tests involving short-run and tightly controlled (e.g., weeded) experiments. Thus, we resampled our 1997 BEF experiment in a restored salt marsh to test for long-term effects of species richness (plantings with 1, 3, and 6 species per 2 x 2 m plot), with multiple ecosystem functions as response variables. Over 11 years, 1- and 6-species assemblages converged on intermediate richness (mean = 3.9 species/ 0.25-m2 plot), and composition changed nonrandomly throughout the site. While three species became rare, the two most productive species became co-dominant. The two dominants controlled and increased shoot biomass, which appeared to decrease species richness. Diversity-function relationships became less positive over 11 years and differed significantly with (a) the species-richness metric (planted vs. measured), and (b) the indicator of function (shoot biomass, height, and canopy layering). The loss of positive relationships between species richness and function in our restored site began soon after we stopped weeding and continued with increasing dominance by productive species. Where species-rich plantings are unlikely to ensure long-term restoration of functions, as in our salt marsh, we recommend dual efforts to establish (1) dominant species that provide high levels of target functions, and (2) subordinate species, which might provide additional functions under current or future conditions.

EMERGING ISSUES FOR THE RESTORATION OF TIDAL MARSH ECOSYSTEMS IN THE CONTEXT OF PREDICTED CLIMATE CHANGE

ABSTRACT There is currently a large regional effort to restore tidal marsh ecosystems in the San Francisco Bay-Delta Estuary involving the commitment of hundreds of millions of dollars and broad landscape-scale habitat manipulations. Although climate change has been on the horizon for many years, recent developments suggest that it must be taken seriously as a factor to be considered in future planning for marsh restoration efforts. Tidal marshes are vulnerable to changes in salinity and inundation rates, both of which will be affected by climate change. Restoration sites may be particularly vulnerable given unpredictable sediment inputs and newly established vegetation. Predicted shifts in snowmelt and altered runoff will change estuarine salinity patterns and could have large-scale impacts on marsh dominance, especially for freshwater marshes. Even relatively small salinity changes could lead to shifts in dominant species, with freshwater marshes being replaced by brackish marshes and brackish marshes converted to salt marsh communities. This will cause a reduction in overall estuarine plant diversity and productivity, with possible reverberations for the estuarine food web. Based on monitoring data from San Francisco Bay marshes, we predict that salinity will have a more immediate impact on tidal marsh vegetation than sea-level rise. However, sea-level rise poses a potentially greater long-term threat, depending on its rate, because the effects of inundation and a more persistent salinity regime could cause widespread marsh loss. If ice sheets in Antarctica and Greenland begin melting at rapid rates, inundation impacts could be catastrophic for coastal marshes. Given the magnitude of these potential changes, we urge the restoration and conservation management community to integrate these contingencies into adaptive management process and to join with the broader community in forging more flexible governance institutions that can respond effectively to large-scale uncertainties and trajectories as they unfold.

Contributions of Organic and Inorganic Matter to Sediment Volume and Accretion in Tidal Wetlands at Steady State

Abstract A mixing model derived from first principles describes the bulk density (BD) of intertidal wetland sediments as a function of loss on ignition (LOI). The model assumes that the bulk volume of sediment equates to the sum of self‐packing volumes of organic and mineral components or BD = 1/[LOI/k1 + (1‐LOI)/k2], where k1 and k2 are the self‐packing densities of the pure organic and inorganic components, respectively. The model explained 78% of the variability in total BD when fitted to 5075 measurements drawn from 33 wetlands distributed around the conterminous United States. The values of k1 and k2 were estimated to be 0.085 ± 0.0007 g cm−3 and 1.99 ± 0.028 g cm−3, respectively. Based on the fitted organic density (k1) and constrained by primary production, the model suggests that the maximum steady state accretion arising from the sequestration of refractory organic matter is ≤ 0.3 cm yr−1. Thus, tidal peatlands are unlikely to indefinitely survive a higher rate of sea‐level rise in the absence of a significant source of mineral sediment. Application of k2 to a mineral sediment load typical of East and eastern Gulf Coast estuaries gives a vertical accretion rate from inorganic sediment of 0.2 cm yr−1. Total steady state accretion is the sum of the parts and therefore should not be greater than 0.5 cm yr−1 under the assumptions of the model. Accretion rates could deviate from this value depending on variation in plant productivity, root:shoot ratio, suspended sediment concentration, sediment‐capture efficiency, and episodic events.