Nitrogen addition does not reduce belowground competition in a salt marsh clonal plant community in Mississippi (USA)

Previous studies have suggested that belowground competition for nutrients influences plant zonation in salt marshes. In this study, I tested the hypothesis that competition for nitrogen structured a clonal plant community in a nitrogen-limited salt marsh in coastal Mississippi, USA. In contrast to most previous field studies that have investigated mechanisms of competition, I examined clonal growth responses of established genets of a nitrogen-demanding low-intertidal species (Spartina alterniflora) to nitrogen addition and the removal of a nitrogen-conserving high-intertidal species (Juncus roemerianus). Nitrogen addition stimulated clonal invasion of the Juncus zone by Spartina but did not reduce the significant competitive effects of Juncus on Spartina. Simulated Juncus shade did not reduce invasion of the Juncus zone by Spartina, indicating that belowground competition reduced clonal invasion. In the last year of the study, the border shifted unexpectedly towards the Spartina zone, resulting in competitive displacement of Spartina by Juncus. Nitrogen addition did not prevent or slow this displacement, further contradicting the nitrogen competition hypothesis. Although growth rates were much more strongly limited by nitrogen in Spartina than in Juncus, nitrogen addition did not cause the displacement of Juncus by Spartina after three growing seasons. I conclude that zonation of Spartina and Juncus is maintained by preemption of space and greater tolerance of low nitrogen supplies by Juncus in the high marsh. These results contrast sharply with findings of reduced belowground competition with nutrient addition in previous studies and highlight the important role of nutrient-mediated competition for space between clonal plants.     

Effect of NaCl on the in vitro Activity of Malate Dehydrogenase in Salt Marsh Halophytes of the U.S.

The in vitro effect of NaCl on NAD-malate dehydrogenase (E.C. 1.1.1.37; MDH) from desalted extracts of roots and leaves of six salt marsh halophytes was investigated. The plants, all native and important constituents of the salt marshes of the east coast of the U.S., included Spartina alterniflora Loisel., Spartina patens (Aiton) Muhl., Distichlis spicata (L.) Greene, Juncus roemerianus Schleele, Salicornia virginica L., and Borrichia frutescens (L.) DC. In the leaf extracts of all species except Borrichia frutescens, the MDH activity was slightly stimulated by NaCl at concentrations around 0.05 M at optimal pH (8.0–8.5) and was reduced by NaCl in higher concentrations. MDH activity in the leaf extract of Borrichia frutescens was more salt-tolerant and maximal activity occurred around 0.25 M NaCl at optimal pH (7.0). Even though similar pH optimums for activity were exhibited in the root and leaf extracts of each species, the MDH activity in the root extract was more salt-tolerant than that in the leaf extract. NaCl at concentrations up to 0.1 M stimulated the MDH activity in the root extracts of all species except that of Borrichia frutescens, which had an optimal activity in 0.5 M NaCl. In the root and leaf extracts of Borrichia frutescens, the activity of cytosol MDH was much more salt-tolerant than that of the mitochondrial MDH. A shift of the optimal pH to more acidic values with increasing concentrations of NaCl was noted in the extracts of all the species except Borrichia frutescens. The action of NaCl on MDH activity appeared to be a general ionic effect as judged by the response of the enzyme activity in the presence of iso-ionic concentrations of other salts and isoosmotic mannitol. Thus, the response of the MDH from five of the salt marsh plants to NaCl is similar to that of glycophytes. However, Borrichia frutescens possesses a salt-tolerant MDH that has optimal activity in a salt concentration as high as that of the environment.     

Identifying high-level salt marshes using a palynomorphic fingerprint with potential implications for tracking sea level change

Palynomorph assemblages from South Carolina salt marshes were analyzed from high marsh, low marsh, and salt panne zones. High marsh zones, dominated by Juncus roemerianus, consistently had a distinctive palynomorphic signature, whereas this was not the case for assemblages from low marsh and salt panne zones.The Juncus environment was distinguishable from all others by the following parameters: 1) higher palynomorph diversity (nearly double the taxonomic diversity present in the other types), 2) the presence of a high percentage (over 10% of the palynomorph count) of Fungal Spore Type A, and 3) the presence of Atrotorquata lineata, a fungal species that was found solely in sediments beneath Juncus marshes. Locating the positions of Juncus environments in cores should allow one to trace the rise and fall of sea level over time.     

EFFECTS OF SULFIDE ON THE GROWTH OF THREE SALT MARSH HALOPHYTES OF THE SOUTHEASTERN UNITED STATES

A hydroponic culture experiment was performed to ascertain whether sediment soluble sulfide at in situ concentrations plays a role in the determination of height forms of Spartina alterniflora in salt marshes of the United States. Additional experiments were conducted for both Spartina cynosuroides and Borrichia frutescens to determine if sulfide also influences the overall distribution of these species in the marsh. In situ soluble sulfide concentrations ranged from 0.02 mm in creek bank sites up to 3.0 mm in the inner marsh. In culture treatments, both plant height and biomass production of S. alterniflora were inhibited at a sulfide concentration as low as 1.0 mm , strongly suggesting a role for sulfide in the determination of height forms in the marsh. Production of S. cynosuroides was inhibited at high sulfide concentrations. However, over a range of concentrations similar to in situ values, no significant reduction in growth was observed, indicating sulfide was not a primary determinant of growth in stands of S. cynosuroides on Sapelo Island, Georgia. A sulfide concentration of 0.5 mm inhibited production in B. frutescens. In situ sulfide concentrations as high as 0.5 mm were found only in mixed stands of Juncus roemerianus and B. frutescens.     

Growth of tissue culture–regenerated salt marsh monocots in a simulated marsh field plot: Implication for wetland creation and restoration

Phenotypically and genetically variable salt marsh plants are needed for wetland creation and restoration efforts. Selected tissue culture regenerants of five salt marsh monocots, Spartina patens, Spartina alterniflora, Juncus gerardi, Juncus roemerianus, and Scirpus robustus, were planted in a simulated marsh field plot that was flood-irrigated with 10 ppt salt water to compare their phenotypic variation for potential use in wetland projects. Plant growth was evaluated after one growing season. Phenotypic variation among regenerants was found in S. alterniflora, S. patens and J. gerardi, indicating the occurrence of somaclonal variation. In S. alterniflora, significant differences occurred among regenerants in stem density. In J. gerardi, significant differences occurred in height and clone circumference. In S. patens, two of the nine regenerants exhibited higher biomass and stem density than some of the other regenerants. By using the random amplified polymorphic DNA (RAPD) technique, genome DNA variation in S. patens regenerants was detected. Genetic variation not only occurred among phenotypically different regenerants, but also among those phenotypically similar for the characteristics measured. Tissue culture–regenerated plants often have desirable genetic characteristics and adaptability as a result of somaclonal variation and may enable a species to perform its ecological functions in created or restored wetlands where ideal environments cannot be achieved. Thus, some previously unrestorable sites may be restorable or marginal marshes made more productive.     

Sulfur storage changed by exotic Spartina alterniflora in coastal saltmarshes of China

For the purpose of ecological engineering, Spartina alterniflora was introduced to China in 1979 and now covers about 112,000 ha of China's coastal lands. It was hypothesized that S. alterniflora could actively change the habitat environment, thus facilitating its competition over native species. In Yancheng Nature Reserve, sulfur storage of sediments and plant tissues was compared among marshes dominated by the exotic S. alterniflora and adjacent native Suaeda salsa and Phragmites australis and bare mudflat. Results showed that the S. alterniflora marsh contained the highest content of water-soluble, adsorbed, carbonate-occluded and total sulfur in the sediment. The sulfur levels were higher in the center than at the edges of the S. alterniflora marsh. Native marshes showed no significant difference in sediment sulfur levels. With greater biomass and higher tissue sulfur concentrations, plant sulfur storage of S. alterniflora vegetation was also larger than those of the native vegetations. Because higher concentrations of sulfur increase the competitive advantage of S. alterniflora over native halophytes, the results of the research showing that S. alterniflora increased marsh sulfur storage may shed light on the mechanism of expansion of monospecific vegetation in coastal China.     

The macrobenthos in Spartina alterniflora salt marshes of the Wanggang tidal-flat,Jiangsu coast,China

On the Jiangsu coast, eastern China, Spartina alterniflora, which was introduced artificially into the region, is becoming a dominant plant species in the inter-tidal salt marshes. In order to evaluate the environmental and ecological impact of the colonization of S. alterniflora, we carried out investigations into the benthic macrofauna of the Spartina marshes of the Wanggang area, central Jiangsu coast, in 2006 and 2007. Based on analysis of the data sets obtained, 12 species of macrobenthos have been identified for the Wanggang salt marsh, including S. alterniflora, Cerithidea cingulata, Littorna scabra, Bithynia fuchsiana, Macrophthalmus japonicus, Uca arcuata, Nereis sp., Boleophthalmus petinirostris, Cyclina sinensis, Bullacta exarata, Angustassiminea castanea and Glaucomya chinensis. The results indicate that some of the native species have adapted to the new ecological environment associated with the cordgrass S. alterniflora. The biomass of macrobenthos varies significantly over different parts of the salt marsh. Further, there is a seasonal change in bio-density, with the density in summer (July and August) > autumn (November) > early summer (May). At the landward edge and over the central part of the S. alterniflora marsh, the bio-diversity is higher than the other areas of the marsh.     

EFFECT OF AN AMMONIUM NITRATE PULSE ON THE GROWTH AND ELEMENTAL COMPOSITION OF NATURAL STANDS OF SPARTINA ALTERNIFLORA AND JUNCUS ROEMERIANUS

A nitrogen (ammonium nitrate) pulse of 200 kg ha“¹ was added to stands of tall (1.0–1.5 m) Spartina alterniflora, short (< 0.5 m) Spartina alterniflora, and Juncus roemerianus in a Georgia salt marsh in July. The major response ten weeks later was an increase in the aerial biomass and a sharp reduction in the C/N ratio in short Spartina alterniflora. One year after the treatment the difference between the biomass in enriched and control plots was greater than ten weeks after treatment, but the C/N ratio in the plants in the treated plots had risen to that of the controls. The availability of nitrogen appears to limit growth in the middle elevation Georgia salt marsh (short S. alterniflora), but not in the lower (tall S. alterniflora) or higher (J. roemerianus) portions.     

Impacts of an alien species (Spartina alterniflora) on the macrobenthos community of Jiangsu coastal inter-tidal ecosystem

Spartina alterniflora, a species vegetating on inter-tidal flats that was introduced from the eastern coast of United States, has become a hot topic, focusing on its invasion within local species in the coastal zone of China. Impacts of S. alterniflora on the inter-tidal macrobenthos community in the Jiangsu coastland are addressed by comparing the macrobenthos characteristics in a mudflat and in a four-year-old Spartina salt marsh that had earlier been a mudflat. During the period October 2002–July 2003, we studied the distribution pattern and diversity of macrobenthos, and discussed their correlation with environmental factors caused by Spartina vegetation. The results showed that a total of 43 macrobenthos species were found, mainly consisting of Mollusca, Crustacea, and Annelida. Ten macrobenthos species were found in the Spartina salt marsh, and 36 species were found in the mudflat. Life forms and functional groups of macrobenthos in the Spartina salt marsh were obviously distrinct from that of the mudflat. The study showed that macrobenthos diversity in the Spartina salt marsh decreased, and the community structure altered obviously, whereas the biomass showed no differences in different seasons. Statistical analysis demonstrated that seasonal change of macrobenthos diversity in the Spartina salt marsh negatively related to content of sediment organic matter, total N, bulk density, height and biomass of Spartina vegetation, and positively related to the density of Spartina. All these differences suggested the obvious effects of the Spartina vegetation on the Jiangsu inter-tidal benthic macroinvertebrate ecology. Furthermore, the investigation also showed that the niche of the native macrobenthos living in the mudflat has been transferred down, seaward, due to the invasion of Spartina in our study site.     

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.     

Modeling the effect of water level on the Nueces Delta marsh community

Water resource development has decreased water delivery to marshes in the Nueces Delta, Corpus Christi, Texas, USA by 45% since 1983, which has led to marsh degradation. Recent management actions will allow for partial hydrological restoration of the marsh, but there is a need to understand the dynamics and the interactive roles of climate and water cycle changes in order to predict changes in salt marshes in the future. In this study, a model of multi-species competition with respect to hydrological change was developed to perform modeling experiments of the effects of water elevation on development of marsh plant species. Nueces Delta plants were divided into two functional groups: (1) clonal stress tolerant plants (Batis maritima, Distichlis spicata, Monanthcloe littoralis, and Salicornia virginica), and (2) clonal dominants (Borrichia frutescens and Spartina alterniflora). Growth rates were calculated for three climate regimes (wet, moderate, and dry), and in three elevation locations (low, mid, and high marsh). The model predicts reductions in plant cover in both drought and moderate conditions. Marsh plant coverage increases only during wet conditions and when there is space available for plant expansion. It is concluded that changes in areal extent of the marsh largely depend on water flow and elevation, which in turn depends on the quantity of fresh water flowing into the marsh. However, under current climate and water management conditions, the marsh will degrade further.     

Formyltetrahydrofolate Synthetase Sequences from Salt Marsh Plant Roots Reveal a Diversity of Acetogenic Bacteria and Other Bacterial Functional Groups

Sixty-two partial formyltetrahydrofolate synthetase (FTHFS) structural gene sequences were recovered from roots of salt marsh plants, including Spartina alterniflora, Salicornia virginica, and Juncus roemerianus. Only S. alterniflora roots yielded sequences grouping with FTHFS sequences from known acetogens. Most other FTHFS or FTHFS-like sequences grouped with those from sulfate-reducing bacteria. Several sequences that grouped with Sphingomonas paucimobilis ligH were also recovered.     

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.     

Effect of the Spartina alterniflora Root-Rhizome System on Salt Marsh Soil Denitrifying Bacteria

Nitrous oxide (N₂O) reductase activity was used as an index of the denitrification potential in salt marsh soils. In a short Spartina alterniflora marsh, the seasonal distribution of N₂O reductase activity indicated a causal relationship between S. alterniflora root-rhizome production and the denitrification potential of the soil system. The relationship was not discerned in samples from a tall S. alterniflora marsh. To further examine the in situ plant-denitrifier interaction in the short S. alterniflora marsh, plots with and without living S. alterniflora were established and analyzed for N₂O reductase activity 5 and 18 months later. In the plots without living Spartina there was a significant reduction in the soil denitrification potential after 18 months, indicating that in the SS marsh the denitrifiers are tightly coupled to the seasonal production of below-ground Spartina macroorganic matter. In plots with intact Spartina, the soil denitrification potential was not altered by NH₄NO₃ or glucose enrichment. However, in plots without living Spartina, there were significant changes in soil N₂O reductase activity, thus indicating that the plants can serve as a “buffer” against this form of pulse perturbation.     

EVALUATION OF PROLINE ACCUMULATION IN THE ADAPTATION OF DIVERSE SPECIES OF MARSH HALOPHYTES TO THE SALINE ENVIRONMENT

Proline accumulation by eight major species of salt marsh halophytes was examined under growth chamber and field conditions. When the plants were exposed to increasing salinities in the growth chamber, they accumulated proline after a threshold salinity had been reached. Three general patterns were apparent. Limonium carolinianum (Walt.) Britt. and Junius roemerianus Scheele began to accumulate proline at 0.25 m NaCl with accumulations up to 63.6 μmoles per gram fresh weight at higher salinities. The C₄ grasses, Spartina alterniflora Loisel., Spartina patens (Aiton) Muhl., and Distichlis spicata (L.) Greene, had threshold salinity levels around 0.5 m NaCl and accumulated proline to 27.4 μmoles per gram fresh weight. The succulents, Salicornia bigelovii Torr., Salicornia virginica L., and Borrichia frutescens (L.) DC, did not accumulate proline until very high salinities (0.7 m) were reached. Water stress imposed by polyethylene glycol instead of NaCl caused similar proline accumulation in the species studied, but to different extents. Field measurements of proline content and soil salinities correlated well with the findings from growth chamber experiments. Rates of proline accumulation and breakdown in L. carolinianum were sufficient for osmotic adjustment by the plant to the changes in interstitial salinity in the marsh. The significance of proline accumulation as an adaptation to the salt marsh environment was species specific. We suggest that proline accumulation is of considerable importance in L. carolinianum and J. roemerianus, important to the C₄ grasses at certain times and in certain locations in the marsh, and of little importance in the succulents.     

Exotic Spartina alterniflora provides compatible habitats for native estuarine crab Sesarma dehaani in the Yangtze River estuary

Although the impact of plant invasions on benthic communities, especially burrowing crabs, has received increasing attention, the results from past studies are mixed. The exotic plant Spartina alterniflora has become the most abundant species in the salt marshes of the Yangtze River estuary since it was first found just over a decade ago, but its effects on crabs in the salt marshes is largely unknown. To examine whether the invasions of this exotic plant affected native crabs, we compared the biomass and abundance of the dominant burrowing crab Sesarma dehaani in an exotic Spartina marsh, native Phragmites australis marsh and mudflats of the Yangtze River estuary, China. To explain the differences of S. dehaani populations between different habitats, feeding preference of S. dehaani for Spartina and Phragmites was investigated. Results showed crab abundance and biomass in the Spartina marsh were significantly greater than those in the Phragmites marsh and mudflats. Soil water content and plant community characteristics in the Spartina marsh also significantly differed in the Phragmites marsh and mudflats. Moreover, the feeding preference experiment showed that crabs consumed Spartina more than twice as much as Phragmites. In summary, this study showed that Spartina provided compatible habitats for native crab S. dehaani through offering suitable food source and moderate environmental conditions.     

Zonation of shrubs in western Atlantic salt marshes

We explored the generality of the processes mediating shrub zonation in western Atlantic salt marshes by comparing the results of our experiments in Georgia, USA with previous studies from Rhode Island, USA. The shrub Borrichia frutescens dominates the terrestrial border of many Georgia salt marshes. Within the shrub zone, physical stress increased at lower elevations, shrubs at lower elevations were stunted, and experimentally reducing physical stress reduced shrub stunting. Below the shrub zone, physical stress increased further, and the grass Spartina alterniflora dominated. Transplant and neighbor-removal experiments indicated that the lower border of the shrub zone was set more by physical stress than by competition, but that the upper border of the grass zone was set primarily by competition with shrubs. Laboratory experiments indicated that S. alterniflora seedlings survived best and shrub seedlings worst in the flooded, salty treatment that mimicked low-marsh conditions. These processes are similar to those maintaining zonation patterns between the shrub Iva frutescens and the rush Juncus gerardi in Rhode Island salt marshes. However, markedly different processes appear to occur further to the north, where woody shrubs are absent from coastal marshes, and further to the south, where woody plants (mangroves) dominate coastal wetlands.     

Physical stress, not biotic interactions, preclude an invasive grass from establishing in forb-dominated salt marshes

Background

Biological invasions have become the focus of considerable concern and ecological research, yet the relative importance of abiotic and biotic factors in controlling the invasibility of habitats to exotic species is not well understood. Spartina species are highly invasive plants in coastal wetlands; however, studies on the factors that control the success or failure of Spartina invasions across multiple habitat types are rare and inconclusive.

Methodology and Principal Findings

We examined the roles of physical stress and plant interactions in mediating the establishment of the smooth cordgrass, Spartina alterniflora, in a variety of coastal habitats in northern China. Field transplant experiments showed that cordgrass can invade mudflats and low estuarine marshes with low salinity and frequent flooding, but cannot survive in salt marshes and high estuarine marshes with hypersaline soils and infrequent flooding. The dominant native plant Suaeda salsa had neither competitive nor facilitative effects on cordgrass. A common garden experiment revealed that cordgrass performed significantly better when flooded every other day than when flooded weekly. These results suggest that physical stress rather than plant interactions limits cordgrass invasions in northern China.

Conclusions and Significance

We conclude that Spartina invasions are likely to be constrained to tidal flats and low estuarine marshes in the Yellow River Delta. Due to harsh physical conditions, salt marshes and high estuarine marshes are unlikely to be invaded. These findings have implications for understanding Spartina invasions in northern China and on other coasts with similar biotic and abiotic environments.     

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     

ELEMENTAL ANALYSIS OF DEPOSITS ON THE ROOTS OF SPARTINA ALTERNIFLORA LOISEL.

Spartina alterniflora, salt marsh cordgrass, is the dominant angiosperm of a large majority of regularly flooded marshes of the Atlantic and Gulf coasts of the United States. In Louisiana, this species often occurs in two distinct zones: a more productive streamside site (adjacent to tidal creeks), and a less productive and sparsely populated inland area. Reddish-brown deposits are present on the roots of streamside Spartina and visually absent from the roots of inland plants. A study of streamside roots using scanning electron microscope and energy dispersive X-ray microanalysis demonstrated that the coatings are restricted to the outer cell wall of the epidermis and are composed primarily of iron. Roots of inland plants have minor iron deposits. Citrate-dithionite extraction of the coatings and subsequent atomic absorption spectrophotometric analysis confirmed these deposits to be iron, although some manganese was present. Approximately 50 times more iron was found on streamside roots compared to roots from inland plants. These results indicate a better developed oxidized rhizosphere associated with streamside Spartina roots than the inland and, hence, the potential for a more favorable environment in which nutrient uptake may proceed.