Salt marsh

salt marsh along the Ōpāwaho / Heathcote River, Christchurch, New Zealand Salt marshes occur on low-energy shorelines in temperate and high-latitudes which can be stable, emerging, or submerging depending if the sedimentation is greater, equal to, or lower than relative sea level rise (subsidence rate plus sea level change), respectively. Commonly these shorelines consist of mud or sand flats (known also as tidal flats or mudflats) which are nourished with sediment from inflowing rivers and streams. These typically include sheltered environments such as embankments, estuaries and the leeward side of barrier islands and spits. In the tropics and sub-tropics they are replaced by mangroves; an area that differs from a salt marsh in that instead of herbaceous plants, they are dominated by salt-tolerant trees. Salt marshes are located among different landforms based on their physical and geomorphological settings. Such marsh landforms include deltaic marshes, estuarine, back-barrier, open coast, embayments and drowned-valley marshes. Deltaic marshes are associated with large rivers where many occur in Southern Europe such as the Camargue, France in the Rhône delta or the Ebro delta in Spain. They are also extensive within the rivers of the Mississippi River Delta in the United States. Such marshes are located in Awhitu Regional Park in Auckland, the Manawatū Estuary, and the Avon Heathcote Estuary / Ihutai in Christchurch. Back-barrier marshes are sensitive to the reshaping of barriers in the landward side of which they have been formed. The low physical energy and high grasses provide a refuge for animals. Many marine fish use salt marshes as nursery grounds for their young before they move to open waters. Birds may raise their young among the high grasses, because the marsh provides both sanctuary from predators and abundant food sources which include fish trapped in pools, insects, shellfish, and worms. == Worldwide occurrence ==

Saltmarshes across 99 countries (essentially worldwide) were mapped by Mcowen et al. 2017. A total of 5,495,089 hectares of mapped saltmarsh across 43 countries and territories are represented in a Geographic Information Systems polygon shapefile. This estimate is at the relatively low end of previous estimates (2.2–40 Mha). A later study conservatively estimated global saltmarsh extent as 90,800 km2 (9,080,000 hectares). The most extensive saltmarshes worldwide are found outside the tropics, notably including the low-lying, ice-free coasts, bays and estuaries of the North Atlantic which are well represented in their global polygon dataset. ==Formation==

The formation begins as tidal flats gain elevation relative to sea level by sediment accretion, and subsequently the rate and duration of tidal flooding decreases so that vegetation can colonize on the exposed surface. The arrival of propagules of pioneer species such as seeds or rhizome portions are combined with the development of suitable conditions for their germination and establishment in the process of colonisation. which can also increase the erosion resistance of the sediments. This assists the process of sediment accretion to allow colonising species (e.g., Salicornia spp.) to grow. These species retain sediment washed in from the rising tide around their stems and leaves and form low muddy mounds which eventually coalesce to form depositional terraces, whose upward growth is aided by a sub-surface root network which binds the sediment. Once vegetation is established on depositional terraces further sediment trapping and accretion can allow rapid upward growth of the marsh surface such that there is an associated rapid decrease in the depth and duration of tidal flooding. As a result, competitive species that prefer higher elevations relative to sea level can inhabit the area and often a succession of plant communities develops. ==Tidal flooding and vegetation zonation==

Coastal salt marshes can be distinguished from terrestrial habitats by the daily tidal flow that occurs and continuously floods the area. Salt Marsh Nature Center in Brooklyn, New York The New England salt marsh is subject to strong tidal influences and shows distinct patterns of zonation. In low marsh areas with high tidal flooding, a monoculture of the smooth cordgrass, Spartina alterniflora dominate, then heading landwards, zones of the salt hay, Spartina patens, black rush, Juncus gerardii and the shrub Iva frutescens are seen respectively. These species all have different tolerances that make the different zones along the marsh best suited for each individual. Plant species diversity is relatively low, since the flora must be tolerant of salt, complete or partial submersion, and anoxic mud substrate. The most common salt marsh plants are glassworts (Salicornia spp.) and the cordgrass (Spartina spp.), which have worldwide distribution. They are often the first plants to take hold in a mudflat and begin its ecological succession into a salt marsh. Their shoots lift the main flow of the tide above the mud surface while their roots spread into the substrate and stabilize the sticky mud and carry oxygen into it so that other plants can establish themselves as well. Plants such as sea lavenders (Limonium spp.), plantains (Plantago spp.), and varied sedges and rushes grow once the mud has been vegetated by the pioneer species. Salt marshes are quite photosynthetically active and are extremely productive habitats. They serve as depositories for a large amount of organic matter and are full of decomposition, which feeds a broad food chain of organisms from bacteria to mammals. Many of the halophytic plants such as cordgrass are not grazed at all by higher animals but die off and decompose to become food for micro-organisms, which in turn become food for fish and birds. ==Sediment trapping, accretion, and the role of tidal creeks==

in Georgia, US The factors and processes that influence the rate and spatial distribution of sediment accretion within the salt marsh are numerous. Sediment deposition can occur when marsh species provide a surface for the sediment to adhere to, followed by deposition onto the marsh surface when the sediment flakes off at low tide. For example, in a study of the Eastern Chongming Island and Jiuduansha Island tidal marshes at the mouth of the Yangtze River, China, the amount of sediment adhering to the species Spartina alterniflora, Phragmites australis, and Scirpus mariqueter decreased with distance from the highest levels of suspended sediment concentrations (found at the marsh edge bordering tidal creeks or the mudflats); decreased with those species at the highest elevations, which experienced the lowest frequency and depth of tidal inundations; and increased with increasing plant biomass. Spartina alterniflora, which had the most sediment adhering to it, may contribute >10% of the total marsh surface sediment accretion by this process. Measured concentrations of suspended sediment in the water column have been shown to decrease from the open water or tidal creeks adjacent to the marsh edge, to the marsh interior, probably as a result of direct settling to the marsh surface by the influence of the marsh canopy. Their typically dendritic and meandering forms provide avenues for the tide to rise and flood the marsh surface, as well as to drain water, Salt marshes do not however require tidal creeks to facilitate sediment flux over their surface Species at higher elevations can benefit from a greater chance of inundation at the highest tides when increased water depths and marsh surface flows can penetrate into the marsh interior. ==Human impacts==

'' (saltmarsh cordgrass). Native to the eastern seaboard of the United States. Considered a noxious weed in the Pacific Northwest The coast is a highly attractive natural feature to humans through its beauty, resources, and accessibility. As of 2002, over half of the world's population was estimated to being living within 60 km of the coastal shoreline, The indirect effects of human activities such as nitrogen loading also play a major role in the salt marsh area. Salt marshes can suffer from dieback in the high marsh and die-off in the low marsh. A study published in 2022 estimates that 22% of saltmarsh loss from 1999–2019 was due to direct human drivers, defined as observable activities occurring at the location of the detected change, such as conversion to aquaculture, agriculture, coastal development, or other physical structures. For centuries, livestock such as sheep and cattle grazed on the highly fertile salt marsh land. Land reclamation for agriculture has resulted in many changes such as shifts in vegetation structure, sedimentation, salinity, water flow, biodiversity loss and high nutrient inputs. There have been many attempts made to eradicate these problems for example, in New Zealand, the cordgrass Spartina anglica was introduced from England into the Manawatū River mouth in 1913 to try and reclaim the estuary land for farming. Terrestrial soils of this nature need to adjust from fresh to saline interstitial water by a change in the chemistry and the structure of the soil, accompanied with fresh deposition of estuarine sediment, before salt marsh vegetation can establish. The vegetation structure, species richness, and plant community composition of salt marshes naturally regenerated on reclaimed agricultural land can be compared to adjacent reference salt marshes to assess the success of marsh regeneration. Upstream agriculture Cultivation of land upstream from the salt marsh can introduce increased silt inputs and raise the rate of primary sediment accretion on the tidal flats, so that pioneer species can spread further onto the flats and grow rapidly upwards out of the level of tidal inundation. As a result, marsh surfaces in this regime may have an extensive cliff at their seaward edge. At the Plum Island estuary, Massachusetts (U.S.), stratigraphic cores revealed that during the 18th and 19th century the marsh prograded over subtidal and mudflat environments to increase in area from 6 km2 to 9 km2 after European settlers deforested the land upstream and increased the rate of sediment supply. Urban development and nitrogen loading '' is a common marine algae found in the salt marsh. The conversion of marshland to upland for agriculture has in the past century been overshadowed by conversion for urban development. Coastal cities worldwide have encroached onto former salt marshes and in the U.S. the growth of cities looked to salt marshes for waste disposal sites. Estuarine pollution from organic, inorganic, and toxic substances from urban development or industrialisation is a worldwide problem The remaining marshes surrounding these urban areas are also under immense pressure from the human population as human-induced nitrogen enrichment enters these habitats. Nitrogen loading through human-use indirectly affects salt marshes causing shifts in vegetation structure and the invasion of non-native species. and with an increasing level of nutrients entering the system from anthropogenic effects, the plant species associated with salt marshes are being restructured through change in competition. This loss in biodiversity is not only seen in flora assemblages but also in many animals such as insects and birds as their habitat and food resources are altered. Sea level rise Due to the melting of Arctic sea ice and thermal expansion of the oceans, as a result of global warming, sea levels have begun to rise. As with all coastlines, this rise in water levels is predicted to negatively affect salt marshes, by flooding and eroding them. While salt marshes are susceptible to threats concerning sea level rise, they are also an extremely dynamic coastal ecosystem. Salt marshes may in fact have the capability to keep pace with a rising sea level, by 2100, mean sea level could see increases between 0.6m to 1.1m. Marshes are susceptible to both erosion and accretion, which play a role in a what is called a bio-geomorphic feedback. Salt marsh vegetation captures sediment to stay in the system which in turn allows for the plants to grow better and thus the plants are better at trapping sediment and accumulate more organic matter. This positive feedback loop potentially allows for salt marsh bed level rates to keep pace with rising sea level rates. Salt marsh vegetation helps to increase sediment settling because it slows current velocities, disrupts turbulent eddies, and helps to dissipate wave energy. Marsh plant species are known for their tolerance to increased salt exposure due to the common inundation of marshlands. These types of plants are called halophytes. Halophytes are a crucial part of salt marsh biodiversity and their potential to adjust to elevated sea levels. With elevated sea levels, salt marsh vegetation would likely be more exposed to more frequent inundation rates and it must be adaptable or tolerant to the consequential increased salinity levels and anaerobic conditions. There is a common elevation (above the sea level) limit for these plants to survive, where anywhere below the optimal line would lead to anoxic soils due to constant submergence and too high above this line would mean harmful soil salinity levels due to the high rate of evapotranspiration as a result of decreased submergence. This lateral accommodation space is often limited by anthropogenic structures such as coastal roads, sea walls and other forms of development of coastal lands. A study by Lisa M. Schile, published in 2014, found that across a range of sea level rise rates, marshlands with high plant productivity were resistant against sea level rises but all reached a pinnacle point where accommodation space was necessary for continued survival. The presence of accommodation space allows for new mid/high habitats to form, and for marshes to escape complete inundation. Mosquito control Earlier in the 20th century, it was believed that draining salt marshes would help reduce mosquito populations, such as Aedes taeniorhynchus, the black salt marsh mosquito. In many locations, particularly in the northeastern United States, residents and local and state agencies dug straight-lined ditches deep into the marsh flats. The end result, however, was a depletion of killifish habitat. The killifish is a mosquito predator, so the loss of habitat actually led to higher mosquito populations, and adversely affected wading birds that preyed on the killifish. These ditches can still be seen, despite some efforts to refill the ditches. ==Crab herbivory and bioturbation==

'' of New Zealand shown here, fill a special niche in salt marsh ecosystems. Increased nitrogen uptake by marsh species into their leaves can prompt greater rates of length-specific leaf growth, and increase the herbivory rates of crabs. The burrowing crab Neohelice granulata frequents SW Atlantic salt marshes where high density populations can be found among populations of the marsh species Spartina densiflora and Sarcocornia perennis. In Mar Chiquita lagoon, north of Mar del Plata, Argentina, Neohelice granulata herbivory increased as a likely response to the increased nutrient value of the leaves of fertilised Spartina densiflora plots, compared to non-fertilised plots. Regardless of whether the plots were fertilised or not, grazing by Neohelice granulata also reduced the length specific leaf growth rates of the leaves in summer, while increasing their length-specific senescence rates. This may have been assisted by the increased fungal effectiveness on the wounds left by the crabs. The salt marshes of Cape Cod, Massachusetts (US), are experiencing creek bank die-offs of Spartina spp. (cordgrass) that has been attributed to herbivory by the crab Sesarma reticulatum. At 12 surveyed Cape Cod salt marsh sites, 10% – 90% of creek banks experienced die-off of cordgrass in association with a highly denuded substrate and high density of crab burrows. Populations of Sesarma reticulatum are increasing, possibly as a result of the degradation of the coastal food web in the region. The bare areas left by the intense grazing of cordgrass by Sesarma reticulatum at Cape Cod are suitable for occupation by another burrowing crab, Uca pugnax, which are not known to consume live macrophytes. The intense bioturbation of salt marsh sediments from this crab's burrowing activity has been shown to dramatically reduce the success of Spartina alterniflora and Suaeda maritima seed germination and established seedling survival, either by burial or exposure of seeds, or uprooting or burial of established seedlings. However, bioturbation by crabs may also have a positive effect. In New Zealand, the tunnelling mud crab Helice crassa has been given the stately name of an 'ecosystem engineer' for its ability to construct new habitats and alter the access of nutrients to other species. Their burrows provide an avenue for the transport of dissolved oxygen in the burrow water through the oxic sediment of the burrow walls and into the surrounding anoxic sediment, which creates the perfect habitat for special nitrogen cycling bacteria. These nitrate reducing (denitrifying) bacteria quickly consume the dissolved oxygen entering into the burrow walls to create the oxic mud layer that is thinner than that at the mud surface. This allows a more direct diffusion path for the export of nitrogen (in the form of gaseous nitrogen (N2)) into the flushing tidal water. ==Microbial life in salt marshes==

The variable salinity, climate, nutrient levels and anaerobic conditions of salt marshes provide strong selective pressures on the microorganisms inhabiting them. In salt marshes, microbes play the main role in nutrient cycling and biogeochemical processing. To date, the microbial community of salt marshes has not been found to change drastically due to human impacts, but the research is still ongoing. Because of the major role of microbes in these environments, it is critical to understand the different processes performed and different microbial players present in salt marshes. Salt marshes provide habitat for chemo(litho)autotrophs, heterotrophs, and photoautotrophs alike. These organisms contribute diverse environmental services such as sulfate reduction, nitrification, decomposition and rhizosphere interactions. Chemo(litho)autotrophs in salt marshes Chemoautotrophs, also known as chemolithoautotrophs, are organisms capable of creating their own energy, from the use of inorganic molecules, and are able to thrive in harsh environments, such as deep sea vents or salt marshes, because they do not depend upon external organic carbon sources for their growth and survival. Some Chemoautotrophic bacterial microorganisms found in salt marshes include Betaproteobacteria and Gammaproteobacteria, both classes including sulfate-reducing bacteria (SRB), sulfur-oxidizing bacteria (SOB), and ammonia-oxidizing bacteria (AOB) which play crucial roles in nutrient cycling and ecosystem functioning. Abundance and diversity of sulfate-reducing chemolithoautotrophs Bacterial chemolithoautotrophs in salt marshes include sulfate-reducing bacteria. In these ecosystems, up to 50% of sedimentary remineralization can be attributed to sulfate reduction. The dominant class of sulfate-reducing bacteria in salt marshes tends to be Deltaproteobacteria. Each type of salt-marsh plant has varying lengths of growing seasons, varying photosynthetic rates, and they all lose varying amounts of organic matter to the ocean, resulting in varying carbon-inputs to the ecosystem. The results from an experiment that was done in a salt marsh in the Yangtze estuary in China, The concentration of reduced sulfur compounds, as well as other possible electron donors, increases with more organic-matter decomposition (by other organisms). Therefore if the ecosystem contains more decomposing organic matter, as with plants with high photosynthetic and littering rates, there will be more electron donors available to the bacteria, and thus more sulfate reduction is possible. As a result, the abundance of sulfate-reducing bacteria increases. The high-photosynthetic-rate, high-litter-rate salt marsh plant, S. alterniflora, was discovered to withstand high sulfur concentrations in the soil, which would normally be somewhat toxic to plants. The abundance of chemolithoautotrophs in salt marshes also varies temporally as a result of being somewhat dependent on the organic C-input from plants in the ecosystem. Since plants grow most throughout the summer, and usually begin to lose biomass around fall during their late stage, the highest input of decomposing organic matter is in the fall. Thus seasonally, the abundance of chemolithotrophs in salt marshes is highest in autumn. The enrichment of nitrates in the water increases denitrification, as well as microbial decomposition and primary productivity. Increases in marsh salinity tend to favor AOB, while higher oxygen levels and lower carbon-to-nitrogen ratios favor AOA. These AOB are important in catalyzing the rate-limiting step within the nitrification process, by using ammonium monooxygenase (AMO), produced from amoA, to convert ammonium (NH4+) into nitrite (NO2-). Specifically, within the class of Betaproteobacteria, Nitrosomonas aestuarii, Nitrosomonas marina, and Nitrosospira ureae are highly prevalent within the salt marsh environment; similarly, within the class of Gammaproteobacteria, Nitrosococcus spp. are key AOB in the marshes. The abundance of these chemolithoautotrophs varies along the salinity gradients present within salt marshes: Nitrosomonas are more prevalent within lower salinity or freshwater regions, while Nitrosospira are found to dominate in higher saline environments. The role of nitrification by AOB in salt marshes critically links ammonia, produced from the mineralization of organic nitrogen compounds, to the process of nitrogen oxidation. Further, nitrogen oxidation is important for the downstream removal of nitrates into nitrogen gas, catalyzed by denitrifiers, from the marsh environment. Cyanobacteria in salt marshes Cyanobacteria are important nitrogen fixers in salt marshes, and provide nitrogen to organisms like diatoms and microalgae. Purple bacteria Oxygen inhibits photosynthesis in purple bacteria, which makes estuaries a favorable habitat for them due to the low oxygen content and high levels of light present, optimizing their photosynthesis. In anoxic environments, like salt marshes, many microbes have to use sulfate as an electron acceptor during cellular respiration instead of oxygen, producing lots of hydrogen sulfide as a byproduct. While hydrogen sulfide is toxic to most organisms, purple bacteria require it to grow and will metabolize it to either sulfate or sulfur, and by doing so allowing other organisms to inhabit the toxic environment. They are very adapted to photosynthesizing in low light environments with bacteriochlorophyll pigments a, c, d, and e, to help them absorb wavelengths of light that other organisms cannot. When co-existing with purple bacteria, they often occupy lower depths as they are less tolerant to oxygen, but more photosynthetically adept. Rhizosphere microbes Fungi Some mycorrhizal fungi, like arbuscular mycorrhiza are widely associated with salt marsh plants and may even help plants grow in salt marsh soil rich in heavy metals by reducing their uptake into the plant, although the exact mechanism has yet to be determined. Bacteria Examining 16S ribosomal DNA found in Yangtze River Estuary, the most common bacteria in the rhizosphere were Proteobacteria such as Betaproteobacteria, Gammaproteobacteria, Deltaproteobacteria, and Epsilonproteobacteria. One such widespread species had a similar ribotype to the animal pathogen S. marcescens, and may be beneficial for plants as the bacteria can break down chitin into available carbon and nitrogen for plants to use. Actinobacteria have also been found in plant rhizosphere in costal salt marshes and help plants grow through helping plants absorb more nutrients and secreting antimicrobial compounds. In Jiangsu, China, Actinobacteria from the suborders Pseudonocardineae, Corynebacterineae, Propionibacterineae, Streptomycineae, Micromonosporineae, Streptosporangineae and Micrococcineae were cultured and isolated from rhizosphere soil. Microbial decomposition activity within salt marshes '' sp. on wheat. This fungus is of the same genus common to salt marsh cordgrass. Another key process among microbial salt marshes is microbial decomposition activity. Nutrient cycling in salt marshes is highly promoted by the resident community of bacteria and fungi involved in remineralizing organic matter. Studies on the decomposition of a salt marsh cordgrass, Spartina alterniflora, have shown that fungal colonization begins the degradation process, which is then finished by the bacterial community. The carbon from Spartina alterniflora is made accessible to the salt marsh food web largely through these bacterial communities which are then consumed by bacterivores. Bacteria are responsible for the degradation of up to 88% of lignocellulotic material in salt marshes. The fungi that make up the decomposition community in salt marshes come from the phylum ascomycota, the two most prevalent species being Phaeosphaeria spartinicola and Mycosphaerella sp. strain 2. ==Restoration and management==

'' spp.) species endemic to the high marsh zone. The perception of bay salt marshes as a coastal 'wasteland' has since changed, acknowledging that they are one of the most biologically productive habitats on earth, rivalling tropical rainforests. Salt marshes are ecologically important, providing habitats for native migratory fish and acting as sheltered feeding and nursery grounds. In the United States and Europe, they are now accorded a high level of protection by the Clean Water Act and the Habitats Directive respectively. With the impacts of this habitats and their importance now realised, a growing interest in restoring salt marshes through managed retreat or the reclamation of land has been established. However, many Asian countries such as China still need to recognise the value of marshlands. With their ever-growing populations and intense development along the coast, the value of salt marshes tends to be ignored and the land continues to be reclaimed. suggests two options available for restoring salt marshes. The first is to abandon all human interference and leave the salt marsh to complete its natural development. These types of restoration projects are often unsuccessful as vegetation tends to struggle to revert to its original structure and the natural tidal cycles are shifted due to land changes. The second option suggested by Bakker et al. (1997) Eventually, the site was bought by the city and the group worked together to restore the area. The project involved removing of invasive species and replanting with native ones, along with public talks to other locals, frequent bird walks and clean-up events. ==Research methods==

There is a diverse range and combination of methodologies employed to understand the hydrological dynamics in salt marshes and their ability to trap and accrete sediment. Sediment traps are often used to measure rates of marsh surface accretion when short term deployments (e.g. less than one month) are required. These circular traps consist of pre-weighed filters that are anchored to the marsh surface, then dried in a laboratory and re-weighed to determine the total deposited sediment. For longer term studies (e.g. more than one year) researchers may prefer to measure sediment accretion with marker horizon plots. Marker horizons consist of a mineral such as feldspar that is buried at a known depth within wetland substrates to record the increase in overlying substrate over long time periods. In order to gauge the amount of sediment suspended in the water column, manual or automated samples of tidal water can be poured through pre-weighed filters in a laboratory then dried to determine the amount of sediment per volume of water. Another method for estimating suspended sediment concentrations is by measuring the turbidity of the water using optical backscatter probes, which can be calibrated against water samples containing a known suspended sediment concentration to establish a regression relationship between the two. Marsh surface elevations may be measured with a stadia rod and transit, electronic theodolite, Real-Time Kinematic Global Positioning System, laser level or electronic distance meter (total station). Hydrological dynamics include water depth, measured automatically with a pressure transducer, or with a marked wooden stake, and water velocity, often using electromagnetic current meters. ==See also==