Interactions among salt marsh plants vary geographically but not latitudinally along the California coast

The strength of species interactions often varies geographically and locally with environmental conditions. Competitive interactions are predicted to be stronger in benign environments while facilitation is expected to be stronger in harsh ones. We tested these ideas with an aboveground neighbor removal experiment at six salt marshes along the California coast. We determined the effect of removals of either the dominant species, Salicornia pacifica, or the subordinate species on plant cover, aboveground biomass and community composition, as well as soil salinity and moisture. We found that S. pacifica consistently competed with the subordinate species and that the strength of competition varied among sites. In contrast with other studies showing that dominant species facilitate subordinates by moderating physical stress, here the subordinate species facilitated S. pacifica shortly after removal treatments were imposed, but the effect disappeared over time. Contrary to expectations based on patterns observed in east coast salt marshes, we did not see patterns in species interactions in relation to latitude, climate, or soil edaphic characteristics. Our results suggest that variation in interactions among salt marsh plants may be influenced by local‐scale site differences such as nutrients more than broad latitudinal gradients.     

Short-term sedimentation in Tagus estuary,Portugal: the influence of salt marsh plants

Short-term sediment deposition was studied at four salt marsh areas in the Tagus estuary. In areas covered with Sarcocornia perennis, Sarcocornia fruticosa, Halimione portulacoides and Spartina maritima and also in the non-vegetated areas, sedimentation was measured as the monthly accumulation of sediments on nylon filters anchored on the soil surface, from August 2000 to May 2001. Our experiments were used also to determine the influence of the different plant species in vertical accretion rates. Short-term sedimentation rates (from 2.8 to 272.3 g m⁻² d⁻¹) did show significant differences when the four salt marshes studied in the Tagus estuary were compared to each others. Salt marshes closer to the sediment sources had higher sedimentation rates. Our results suggest that the salt marsh type and surface cover may provide small-scale variations in sedimentation and also that sediment deposition values do change according to the position of the different plant species within the salt marsh. Sedimentation is an essential factor in salt marsh vertical accretion studies and our investigation may provide support to help forecast the adaptative response of the Tagus estuary wetlands to future sea level rise.     

Does low temperature prevent <Emphasis Type="Italic">Spartina alterniflora</Emphasis> from expanding toward the austral-most salt marshes?

Along the Atlantic coast of South America, the northern salt marshes (lower than 43°S) are dominated by Spartina species while the southern salt marshes (greater than 43°S) are dominated by Sarcocornia perennis. The most abundant Spartina species are Spartina densiflora which is present in most coastal marshes, and Spartina alterniflora that was never recorded above the ~42°25′S. It is not clear why S. alterniflora has not succeeded in the southern marshes, in which the low marsh zone remains as an extensive bared mud flat. We address the hypothesis that the absence of S. alterniflora in the south is driven by the cold temperatures inversely related with increasing latitudes along the East coast of Patagonia. To evaluate this hypothesis, we carried out an experiment in which we manipulated the temperature in combination with frost formation and photoperiod. We found that cold temperature produced a negative effect on S. alterniflora, and this effect seems accentuated by the frost but not by the reduction in the photoperiod. Our results support the hypothesis that the absence of S. alterniflora in the southernmost salt marshes of Patagonia is a consequence of the frost as an outcome of the co-occurrence of low temperature and high humidity. The importance of our results are discussed in the context of the global warming and how Spartina species enlarge their distributional range toward higher latitudes.     

Ecological processes shaping Central Patagonian salt marsh landscapes

Plant zonation is one of the most conspicuous ecological features of salt marshes worldwide. In this work we used a combination of field transplant and greenhouse experiments to evaluate the importance of interspecific interactions and physical stress in the determination of the major plant zonation patterns in Central Patagonian salt marshes. There, Spartina alterniflora dominates the low marsh, and Sarcocornia perennis the high marsh. We addressed two questions: (i) What prevents Spartina alterniflora from colonizing the Sarcocornia perennis‐dominated high marsh zone? and (ii) What prevents Sarcocornia perennis from colonizing the Spartina alterniflora‐dominated low marsh zone? Our experimental transplants combined with neighbour exclusion treatments showed that the presence of Sarcocornia perennis negatively affects Spartina alterniflora, preventing it from surviving and/or spreading. Complementary field transplant and greenhouse experiments showed that Sarcocornia perennis did not survive the frequent tidal submersion by approximately 1.5 m of turbid seawater in the Spartina alterniflora zone, but its survival was independent of the presence of Spartina neighbours, and of the strong soil anoxia as well. Our results suggest that Spartina alterniflora is excluded by Sarcocornia perennis towards the low marsh, where frequent and prolonged submersion limit the survival of the latter. We provide and discuss key baseline information to facilitate the future design of ecophysiological experiments designed to accurately identify the exact mechanisms acting in every situation.     

Aboveground competition influences density‐dependent effects of cordgrass on sediment biogeochemistry

Interspecific interactions between plants influence plant phenotype, distribution, abundance, and community structure. Each of these can, in turn, impact sediment biogeochemistry. Although the population and community level impacts of these interactions have been extensively studied, less is known about their effect on sediment biogeochemistry. This is surprising given that many plants are categorized as foundation species that exert strong control on community structure. In southern California salt marshes, we used clipping experiments to manipulate aboveground neighbor presence to study interactions between two dominant plants, Pacific cordgrass (Spartina foliosa) and perennial pickleweed (Sarcocornia pacifica). We also measured how changes in cordgrass stem density influenced sediment biogeochemistry. Pickleweed suppressed cordgrass stem density but had no effect on aboveground biomass. For every cordgrass stem lost per square meter, porewater ammonium increased 0.3–1.0 µM. Thus, aboveground competition with pickleweed weakened the effects of cordgrass on sediment biogeochemistry. Predictions about plant–soil feedbacks, especially under future climate scenarios, will be improved when plant–plant interactions are considered, particularly those containing dominant and foundation species.     

Three new species of <Emphasis Type="Italic">Sarcocornia</Emphasis> (Chenopodiaceae) from South Africa

Sarcocornia decussata, S. freitagii and S. tegetaria from South Africa are described as new taxa. Sarcocornia decussata and S. freitagii are narrow endemics of the West Coast of South Africa where they are found in inland saline habitats (quartz patches, salt pans and saline alluvia) while S. tegetaria is an endemic of southern African coasts (spanning Namibia, South Africa and Mozambique) where it is confined to low-lying intertidal habitats of estuaries.     

The influence of rainfall variability on the species composition of a northern California salt marsh plant assemblage

Natural variability in the species composition of salt marsh plant assemblage was studied for 4 years (1986–90) at Bolinas Lagoon, California (USA). The study was conducted during a period in which little or no physical disturbance occurred in the marsh. During the study, Bolinas Lagoon experienced highly variable rainfall, both within and between years. In years with average or below average winter and spring rainfall, the cover of Salicornia virginica increased relative to that of the other species. During the one year with much higher than average spring rainfall, the cover of Salicornia declined and rare species increased in relative abundance. These patterns suggest that under typical marsh conditions, the plant assemblage experiences stress due to low availability of freshwater and high soil salinity. Under such conditions, the stress-tolerant Salicornia dominates the assemblage. If, however, there is abundant rainfall early in the growing season, the other species are released from stress and are able to increase in cover. The marsh of Bolinas Lagoon appears to be resistant to change and able to tolerate stress. Few studies have examined natural variability in species in undisturbed marshes and more of such studies must be made in order to understand whether or not marshes are in general resistant to change caused by rainfall variability.Abbreviations MLLW = mean lower low water     

Ascomycete fungal communities associated with early decaying leaves of Spartina spp. from central California estuaries

Ascomycetous fungi play an important role in the early stages of decomposition of Spartina alterniflora, but their role in the decomposition of other Spartina species has not been investigated. Here we use fingerprint (terminal restriction fragment length polymorphism) and phylogenetic analyses of the 18S to 28S internal transcribed spacer region to compare the composition of the ascomycete fungal communities on early decay blades of Spartina species (Spartina alterniflora, Spartina densiflora, Spartina foliosa, and a hybrid (S. alterniflora × S. foliosa)) collected from three salt marshes in San Francisco Bay and one in Tomales Bay, California, USA. Phaeosphaeria spartinicola was found on all samples collected and was often dominant. Two other ascomycetes, Phaeosphaeria halima and Mycosphaerella sp. strain 2, were also common. These three species are the same ascomycetes previously identified as the dominant fungal decomposers on S. alterniflora on the east coast. Ascomycetes appeared to exhibit varying degrees of host specificity, demonstrated by grouping patterns on phylogenetic trees. Neither the exotic S. alterniflora nor the hybrid supported fungal flora different from that of the native S. foliosa. However, S. densiflora had a significantly different fungal community than the other species, and hosted at least two unique ascomycetes. Significant differences in the fungal decomposer communities were also detected within species (two clones of S. foliosa), but these were minor and may be due to morphological differences among the plants.     

CULTURE STUDIES ON THE RELATIONSHIP BETWEEN SCHIZYMENIA AND HAEMATOCELIS (GIGARTINALES,RHODOPHYCEAE) FROM THE PACIFIC COAST OF NORTH AMERICA1

Carpospores from Schizymenia pacifica (Kylin) Kylin (Gymnophlaeaceae) from California formed crusts anatomically identical to Haematocelis rubens J. Agardh (Cruoriaceae). Tetraspores of H. rubens from Monterey, California, and Baja California, Mexico, germinated to form basal discs from which arose upright multiaxial blades with a filamentous medulla and cortical gland cells. Pro-carps and spermatangia were present on the same blades; subsequently, cystocarps characteristic of Schizymenia pacifica developed. Re-examination of herbarium specimens suggests that the foliose tetrasporangial phases previously reported as S. pacifica are referable to Halymenia, Dilsea, Cryptonemia, or Turnerella. Schizymenia pacifica (type locality: San Juan Islands, Washington) thus is considered to be the gametophyte in the life history of Haematocelis rubens (type locality: Brest, France), which has also been reported to be the tetrasporophyte of S. dubyi (Chauvin ex Duby) J. Agardh (type locality: Cherbourg, France). Atlantic and Pacific gametophytes and tetrasporophytes are anatomically very similar, suggesting that only one species is involved, but critical studies must be made before a decision on this taxonomic question can be reached. Haematocelis zonalis Dawson et Neushul (type locality: Anacapa Island, California) is considered to be a growth form of the tetrasporangial phase of S. pacifica.     

Changes in pelagic Polychaete assemblages along the California current system

The composition, abundance, species richness and structural changes of the planktonic polychaete assemblages were analysed along a latitudinal transect in the California Current System (California, U.S.A. and Baja California, Mexico). The biological (species and abundance) and physical (temperature and salinity) data were analysed using Principal Component Analysis (PCA). The principal water masses in the survey area were determined. Twenty-four holoplanktonic species belonging to families Alciopidae, Iospilidae, Lopadorhynchidae, Tomopteridae and Typhloscolecidae were identified. Three clear species assemblages were discerned in the PCA results: 1. A `north group' (from Oregon-California border to San Francisco), with relatively high species richness (11) and the highest mean abundance (121 ind. per 500 m<sup>3</sup>) was characterised by Tomopteris septentrionalis, T. planktonis, Plotohelmis tenuis, and Travisiopis lobifera. California Current Water and Subtropical Central Water were present in the area occupied by this assemblage. 2. A `south group' (from off Bahía Magdalena to Cabo San Lucas), with the highest species richness (16), but low mean abundance (37.8 ind. per 500 m³); it included tropical affinity species, such Lopadorhynchus henseni, Tomopteris nationalis, and Travisiopsis dubia. In concordance Surface Equatorial Water was identified in this region. 3. A `transition group' (between the north and south regions) recorded the lowest mean abundance (2.3 ind. per 500 m³) and species richness (9). Only the California Current Water was detected in this area. The spatial pattern of species richness found along of this transect, was at least, partially due to the planktonic productivity distribution in the epipelagic region and the influence of several water masses coming from different directions.     

鄂尔多斯台地盐沼滩地微生物群落与土壤条件分析

[背景]我国北方内陆区与平原区土地盐碱化问题严重,针对微生物如何在极端盐碱地植被演替过程中发挥作用的研究鲜有报道.[目的]研究鄂尔多斯台地5种不同植被类型的盐沼湿地微生物群落与土壤条件的关系,筛选出耐盐碱菌群及影响耐盐碱菌群的土壤环境因子.[方法]采用高通量测序技术,对其微生物细菌群落组成进行了比对分析.[结果]鄂尔多...     

Biomass and aboveground production of four angiosperms in Cantabrian (N. Spain) salt marshes

New data of aboveground biomass and production of four angiosperms over a 12 month period for the Cantabrian Sea salt marshes (Bay of Biscay, N. Spain) are presented. Based on harvest methods, maximum aboveground total biomass values for Spartina maritima (Curtis) Fernald, Spartina alterniflora Loisel, Salicornia ramosissima J. Woods and Halimione portulacoides (L.) Aellen were 628, 1109, 480 and 1267 gm^-2, respectively. We conclude that although a slight latitudinal gradient in biomass is revealed in the data compiled with reference to some of the species studied, more work is neccesary in order to assess the potential productivity of these ecosystems on the coasts of Europe and/or to make comparisons with salt marshes of the American coasts. Annual net aerial primary production estimates using Smalley's method were: 296, 1160, 486 and 952 gm^-2yr^-1, for Spartina maritima, Spartina alterniflora, Salicornia ramosissima and Halimione portulacoides, respectively. These results together with turnover rate estimates point to the lack of vigour of the native S. maritima, while the exotic S. alterniflora, which seems to be spreading along the Cantabrian estuaries, behaves like a veritable pionner throughout the low marshes in this region.     

A new species of Sanicula L. (Umbelliferae/Apiaceae) endemic to Baja California, Mexico

A new species of the genusSanicula (sect.Sanicoria) endemic to the southern Sierra Juárez of Baja California, Mexico, is described and illustrated.Sanicula moranii resemblesS. deserticola andS. bipinnatifida, but has thicker basal leaves, with broader petioles and rachises, and shorter fruit prickles that are confined to the apical part of the mericarps. The taxonomic relationships of these species are discussed, along with aspects of their distribution and habitat.

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Resumen  Una neuva especie del géneroSanicula (secciónSanicoria) endémica de la Sierra Juárez de Baja California, México, es descrita e ilustrada.Sanicula moranii presenta similitud conS. deserticola yS. bipinnatifida, no obstante tiene gruesas hojas basales con pecíolos y raquis anchos, y espínulas del fruto más cortas que se distribuyen únicamente en la parte apical de los mericarpos. Además, se comentan las relaciones taxonómicas de las tres especies, así como algunos aspectos de sus distribuciones y hábitats.

     

Photosynthetic responses to light intensity of <Emphasis Type="Italic">Sarcocornia</Emphasis> taxa (Chenopodiaceae)

Salt marshes show a characteristic zonation of species distribution, which is correlated with marsh elevation. Radiation intensity and photoperiod change throughout the tidal frame. Photosynthetic response to light regime, 90–2450 μmol/(m² s), was determined in the laboratory for four closely related halophytic taxa of the genus Sarcocornia (Chenopodiaceae), which inhabit different positions in the tidal frame. Sarcocornia fruticosa, which germinates below vegetation cover and is found at high levels in the tidal frame, had the lowest maximum net photosynthetic rate and stomata conductance values. The two S. perennis subspecies demonstrated intermediate maximum net photosynthetic rates but S. perennis ssp. alpini reached light saturation point at higher light intensities. S. perennis ssp. perennis, found in the lowest elevations of the marshes, spends a significant proportion of its time submerged and therefore needs to take full advantage of available light. S. perennis ssp. alpini is exposed to very high light intensities in open salt pans at high elevations. The hybrid, S. perennis x fruticosa, which is currently found at intermediate elevations with less frequent inundation, had the highest net photosynthetic rate and chlorophyll a content. The ability to cope with high light levels may help to explain one of the environmental parameters, which affects distribution of four taxa throughout the tidal frame and also raise intriguing questions about the future role of the hybrid in the successive development of these marsh systems.     

High site fidelity and low site connectivity in temperate salt marsh fish populations: a stable isotope approach

Adult and juvenile fish utilise salt marshes for food and shelter at high tide, moving into adjacent sublittoral regions during low tide. Understanding whether there are high levels of site fidelity for different species of coastal fish has important implications for habitat conservation and the design of marine protected areas. We hypothesised that common salt marsh fish species would demonstrate a high site fidelity, resulting in minimal inter-marsh connectivity. Carbon (¹³C) and nitrogen (¹⁵N) stable isotope ratios of larvae and juveniles of five common salt marsh fish (Atherina presbyter, Chelon labrosus, Clupea harengus, Dicentrarchus labrax, Pomatoschistus microps), seven types of primary producer and seven secondary consumer food sources were sampled in five salt marshes within two estuary complexes along the coast of south-east England. Significant differences in ¹³C and ¹⁵N signatures between salt marshes indicated distinct sub-populations utilising the area of estuary around each salt marsh, and limited connectivity, even within the same estuary complex. ¹⁵N ratios were responsible for the majority of inter-marsh differences for each species and showed similar site-specific patterns in ratios in primary producers, secondary consumers and fish. Fish diets (derived from isotope mixing models) varied between species but were mostly consistent between marsh sites, indicating that dietary shifts were not the source of variability of the inter-marsh isotopic signatures within species. These results demonstrate that for some common coastal fish species, high levels of site fidelity result in individual salt marshes operating as discrete habitats for fish assemblages.     

Food web analysis of southern California coastal wetlands using multiple stable isotopes

Carbon, nitrogen, and sulfur stable isotopes were used to characterize the food webs (i.e., sources of carbon and trophic status of consumers) in Tijuana Estuary and San Dieguito Lagoon. Producer groups were most clearly differentiated by carbon, then by sulfur, and least clearly by nitrogen isotope measurements. Consumer ¹⁵N isotopic enrichment suggested that there are four trophic levels in the Tijuana Estuary food web and three in San Dieguito Lagoon. A significant difference in multiple isotope ratio distributions of fishes between wetlands suggested that the food web of San Dieguito Lagoon is less complex than that of Tijuana Estuary. Associations among sources and consumers indicated that inputs from intertidal macroalgae, marsh microalgae, and Spartina foliosa provide the organic matter that supports invertebrates, fishes, and the light-footed clapper rail (Rallus longirostris levipes). These three producers occupy tidal channels, low salt marsh, and mid salt marsh habitats. The only consumer sampled that appears dependent upon primary productivity from high salt marsh habitat is the sora (Porzana carolina). Two- and three-source mixing models identified Spartina as the major organic matter source for fishes, and macroalgae for invertebrates and the light-footed clapper rail in Tijuana Estuary. In San Dieguito Lagoon, a system lacking Spartina, inputs of macroalgae and microalgae support fishes. Salicornia virginica, S. subterminalis, Monanthochloe littoralis, sewage- derived organic matter, and suspended particulate organic matter were deductively excluded as dominant, direct influences on the food web. The demonstration of a salt marsh–channel linkage in these systems affirms that these habitats should be managed as a single ecosystem and that the restoration of intertidal marshes for endangered birds and other biota is compatible with enhancement of coastal fish populations; heretofore, these have been considered to be competing objectives. Received: 24 April 1996 / Accepted: 24 October 1996     

Co-development of wetland soils and benthic invertebrate communities following salt marsh creation

The development of wetland soil characteristics andbenthic invertebrate communities were evaluated increated Spartina alterniflorasalt marshes inNorth Carolina ranging in age from 1 to 25 years-old.A combination of measurements from different-agecreated marshes as well as periodic measurements overtime on two marshes were used to (1) document rates ofwetland pedogenesis, especially soil organic matter,and, (2) explore relationships between soil andbenthic invertebrate community development. Soilmacro-organic matter (MOM, the living and dead rootand rhizome mat), organic C and N increased and bulkdensity decreased during the 25 years following marshestablishment. The most dramatic changes in bulkdensity, MOM, C and N occurred within the upper 10 cmof the soil with lesser changes below this depth.Created marshes were sinks for organic C (90–140g·m^-2·yr^-1) and N (7–11g·m^-2·yr^-1) but not for P (0–1g·m^-2·yr^-1). The density of benthicinvertebrates (>250 m) and subsurface-depositfeeding oligochaetes also increased over time oncreated salt marshes. Invertebrate and oligochaetedensity were strongly related to MOM content(r²= 0.83–0.87) and soil organic C(r²= 0.52–0.82) and N (r²= 0.62–0.84). Thesefindings suggest that, in created salt marshes,development of the benthic invertebrate community istied to marsh soil formation, especially accumulationof organic matter as MOM and soil. Field studies thatmanipulate the quantity and quality of soil organicmatter are needed to elucidate the relationshipbetween salt marsh pedogenesis and benthicinvertebrate community development.     

Decomposition of Sarcocornia quinqueflora on an Iron-Smelting Slag Substrate

Restoration of salt marsh habitat is becoming more common in Australia. However, little is known about restoring salt marshes on substrates contaminated by slag from iron smelting, which could affect microbial activity. This study, conducted near Newcastle, Australia, compares initial C, N, and P mass and decomposition of Sarcocornia quinqueflora (glasswort or samphire) from (1) a restoration site with a slag‐and‐mud substrate, (2) the restoration site's donor marsh, and (3) other nearby sites sampled to provide information on background variability. A litterbag technique with a 180‐day incubation period was used to quantify total, C, N, and P mass losses from decomposition. Although there were significant differences between sites in initial N mass and loss of C and P over the period of our study, the presence of slag did not slow decomposition rates as measured using litterbags. Further work is needed to assess other aspects of wetland structure and function on slag substrates.     

Is the salt marsh vegetation a determining factor in the sedimentation processes?

Many authors have referred to the important role of vegetation in the consolidation of salt marsh sediments, but experiments previously carried out by us have shown results that do not always agree with these statements. In other words, the type of salt marsh surface coverage is not the main factor that contributes to the consolidation of sediments. To test this hypothesis different Portuguese salt marsh stations (species/unvegetated areas) from two sites, Tagus estuary (Corroios and Pancas) and Ria de Aveiro (Barra and Verdemilho), were compared to evaluate their influence on suspended matter deposition on the salt marsh surface. A short-term sedimentation study was performed within stands of Spartina maritima, Halimione portulacoides, Sarcocornia perennis subsp. perennis and unvegetated areas, by analysing the deposition of sediment material on nylon filters anchored to the marsh surface. Numerical results obtained from hydrodynamic models coupled to a Lagrangean module implemented for the Ria de Aveiro and the Tagus Estuary, namely the root-mean square velocity (V _rms) and residual velocity of tides, were also used. Average sedimentation rates (mean value between the different surface cover in a salt marsh) showed a seasonal trend more or less defined but with significantly different values between sites and salt marshes. Sedimentation rates varied between marshes: there are significant differences between Pancas and the other three marshes, but only significant differences in sedimentation rates between Spartina and Sarcocornia. Despite the important role of vegetation in the consolidation of salt marsh sediments, our results suggest that, the position of stations and related abiotic conditions in the salt marshes are determining factors of variation to take into account in the studies related with the stabilization and survival of salt marshes facing sea level rise. Handling editor: P. Viaroli     

Nitrogen further promotes a dominant salt marsh plant in an increasingly saline environment

Aims Human alterations of the environment are combining in unprecedented ways, making predictions of alterations to natural communities a difficult and pressing challenge. Estuarine systems have been subject to a high degree of modification, including increased nitrogen (N) inputs and altered salinity, factors important in shaping estuarine plant communities. As human populations increase and the climate changes, both N and salinity levels are likely to increase in these coastal marshes. Our objective was to evaluate the interactive effects of N and salinity on US West Coast salt marsh species; in particular, the performance of the dominant species Sarcocornia pacifica (pickleweed) alone and in mixed species assemblages. We expected increased salinity to favor S. pacifica but that N enrichment could help maintain greater species richness through use of N in salinity tolerance mechanisms.Methods We crossed treatments of N (added or not) and salinity (salt added or not) in a field experiment at a salt marsh in the San Francisco Estuary, California, USA, in each of three habitats: (i) monotypic pickleweed on the marsh plain, (ii) monotypic pickleweed along channels and (iii) mixed assemblages along channels. In a greenhouse experiment, we crossed treatments of N (added or not) and salinity (at three levels to simulate brackish to saline conditions) in (i) pots of pickleweed only and (ii) the same species mix as in the field.Important findings N addition doubled S. pacifica biomass and branching in both channel and marsh plain habitats regardless of salinity and greatly increased its dominance over Distichlis spicata and Jaumea carnosa in mixed assemblages along channels. In the greenhouse, S. pacifica biomass increased 6- to 10-fold with N addition over the range of salinities, while D. spicata and J. carnosa biomass increased with N addition only at lower salinity levels. Thus, while localized management could influence outcomes, expected overall increases in both N and salinity with human population growth and climate change are likely to enhance the production of S. pacifica in US West Coast marshes while reducing the diversity of mixed species assemblages. This decline in diversity may have implications for the resilience of marshes already subject to multiple stressors as the climate changes.