A salt marsh, saltmarsh or salting, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastalintertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. It is dominated by dense stands of salt-tolerant plants such as herbs, grasses, or low shrubs.[1][2] These plants are terrestrial in origin and are essential to the stability of the salt marsh in trapping and binding sediments. Salt marshes play a large role in the aquatic food web and the delivery of nutrients to coastal waters. They also support terrestrial animals and provide coastal protection.[2]
Salt marshes have historically been endangered by poorly implemented coastal management practices, with land reclaimed for human uses or polluted by upstream agriculture or other industrial coastal uses. Additionally, sea level rise caused by climate change is endangering other marshes, through erosion and submersion of otherwise tidal marshes.[3][4] However, recent acknowledgment by both environmentalists and larger society for the importance of saltwater marshes for biodiversity, ecological productivity and other ecosystem services, such as carbon sequestration, have led to an increase in salt marsh restoration and management since the 1980s.
Salt marshes occur on low-energy shorelines in temperate and high-latitudes[5] 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 abbreviated to mudflats) which are nourished with sediment from inflowing rivers and streams.[6] 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.[1]
Most salt marshes have a low topography with low elevations but a vast wide area, making them hugely popular for human populations.[7] 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 Delta in the United States.[2] In New Zealand, most salt marshes occur at the head of estuaries in areas where there is little wave action and high sedimentation.[8] 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.[2] They are common along much of the eastern coast of the United States and the Frisian Islands. Large, shallow coastal embayments can hold salt marshes with examples including Morecambe Bay and Portsmouth in Britain and the Bay of Fundy in North America.[2]
Salt marshes are sometimes included in lagoons, and the difference is not very marked; the Venetian Lagoon in Italy, for example, is made up of these sorts of animals and or living organisms belonging to this ecosystem. They have a big impact on the biodiversity of the area. Salt marsh ecology involves complex food webs which include primary producers (vascular plants, macroalgae, diatoms, epiphytes, and phytoplankton), primary consumers (zooplankton, macrozoa, molluscs, insects), and secondary consumers.[9]
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.[10]
Worldwide occurrence
Saltmarshes across 99 countries (essentially worldwide) were mapped by Mcowen et al. 2017.[11] 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 90,800km2 (9,080,000 hectares).[12] 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.[11]
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.[13] 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.[14] When rivers and streams arrive at the low gradient of the tidal flats, the discharge rate reduces and suspended sediment settles onto the tidal flat surface, helped by the backwater effect of the rising tide.[6] Mats of filamentous blue-green algae can fix silt and clay sized sediment particles to their sticky sheaths on contact[15] which can also increase the erosion resistance of the sediments.[16] 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.[17] 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.[13]
Coastal salt marshes can be distinguished from terrestrial habitats by the daily tidal flow that occurs and continuously floods the area.[1] It is an important process in delivering sediments, nutrients and plant water supply to the marsh.[7] At higher elevations in the upper marsh zone, there is much less tidal inflow, resulting in lower salinity levels.[1]Soil salinity in the lower marsh zone is fairly constant due to everyday annual tidal flow. However, in the upper marsh, variability in salinity is shown as a result of less frequent flooding and climate variations. Rainfall can reduce salinity and evapotranspiration can increase levels during dry periods.[1] As a result, there are microhabitats populated by different species of flora and fauna dependent on their physiological abilities. The flora of a salt marsh is differentiated into levels according to the plants' individual tolerance of salinity and water table levels. Vegetation found at the water must be able to survive high salt concentrations, periodical submersion, and a certain amount of water movement, while plants further inland in the marsh can sometimes experience dry, low-nutrient conditions. It has been found that the upper marsh zones limit species through competition and the lack of habitat protection, while lower marsh zones are determined through the ability of plants to tolerate physiological stresses such as salinity, water submergence and low oxygen levels.[18][19]
High marsh in the Marine Park 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.[19] 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.[18] 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
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.[13] The amount of sediment adhering to salt marsh species is dependent on the type of marsh species, the proximity of the species to the sediment supply, the amount of plant biomass, and the elevation of the species.[20] 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.[20]
Salt marsh species also facilitate sediment accretion by decreasing current velocities and encouraging sediment to settle out of suspension.[13] Current velocities can be reduced as the stems of tall marsh species induce hydraulic drag, with the effect of minimising re-suspension of sediment and encouraging deposition.[21] 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,[20][21][22] probably as a result of direct settling to the marsh surface by the influence of the marsh canopy.[21][22]
Inundation and sediment deposition on the marsh surface is also assisted by tidal creeks[22] which are a common feature of salt marshes.[6][13][17][22][23] Their typically dendritic and meandering forms provide avenues for the tide to rise and flood the marsh surface, as well as to drain water,[17] and they may facilitate higher amounts of sediment deposition than salt marsh bordering open ocean.[23] Sediment deposition is correlated with sediment size: coarser sediments will deposit at higher elevations (closer to the creek) than finer sediments (further from the creek). Sediment size is also often correlated with particular trace metals, and thus tidal creeks can affect metal distributions and concentrations in salt marshes, in turn affecting the biota.[24] Salt marshes do not however require tidal creeks to facilitate sediment flux over their surface[21] although salt marshes with this morphology seem to be rarely studied.
The elevation of marsh species is important; those species at lower elevations experience longer and more frequent tidal floods and therefore have the opportunity for more sediment deposition to occur.[20][25] 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.[22]
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 60km of the coastal shoreline,[2] making coastlines highly vulnerable to human impacts from daily activities that put pressure on these surrounding natural environments. In the past, salt marshes were perceived as coastal 'wastelands,' causing considerable loss and change of these ecosystems through land reclamation for agriculture, urban development, salt production and recreation.[7][26][27] 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.[12] Additionally, 30% of saltmarsh gain over this same time period were also due to direct drivers, such as restoration activities or coastal modifications to promote tidal exchange.[12]
Land reclamation
Reclamation of land for agriculture by converting marshland to upland was historically a common practice.[7]Dikes were often built to allow for this shift in land change and to provide flood protection further inland. In recent times intertidal flats have also been reclaimed.[28] For centuries, livestock such as sheep and cattle grazed on the highly fertile salt marsh land.[1][29] 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 Manawatu River mouth in 1913 to try and reclaim the estuary land for farming.[8] A shift in structure from bare tidal flat to pastureland resulted from increased sedimentation and the cordgrass extended out into other estuaries around New Zealand. Native plants and animals struggled to survive as non-natives out competed them. Efforts are now being made to remove these cordgrass species, as the damages are slowly being recognized.
In the Blyth estuary in Suffolk in eastern England, the mid-estuary reclamations (Angel and Bulcamp marshes) that were abandoned in the 1940s have been replaced by tidal flats with compacted soils from agricultural use overlain with a thin veneer of mud. Little vegetation colonisation has occurred in the last 60–75 years and has been attributed to a combination of surface elevations too low for pioneer species to develop, and poor drainage from the compacted agricultural soils acting as an aquiclude.[30] 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.[14] 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.[31]
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.[32] 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 6km2 to 9km2 after European settlers deforested the land upstream and increased the rate of sediment supply.[33]
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[28] and the sediment in salt marshes may entrain this pollution with toxic effects on floral and faunal species.[32] Urban development of salt marshes has slowed since about 1970 owing to growing awareness by environmental groups that they provide beneficial ecosystem services.[7] They are highly productive ecosystems, and when net productivity is measured in g m−2 yr−1 they are equalled only by tropical rainforests.[28] Additionally, they can help reduce wave erosion on sea walls designed to protect low-lying areas of land from wave erosion.[14]
De-naturalisation of the landward boundaries of salt marshes from urban or industrial encroachment can have negative effects. In the Avon-Heathcote estuary/Ihutai, New Zealand, species abundance and the physical properties of the surrounding margins were strongly linked, and the majority of salt marsh was found to be living along areas with natural margins in the Avon / Ōtākaro and Ōpāwaho / Heathcote river outlets; conversely, artificial margins contained little marsh vegetation and restricted landward retreat.[34] 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.[18]
Human impacts such as sewage, urban run-off, agricultural and industrial wastes are running into the marshes from nearby sources. Salt marshes are nitrogen limited[18][35] 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.[7] For example, the New England salt marsh is experiencing a shift in vegetation structure where S. alterniflora is spreading from the lower marsh where it predominately resides up into the upper marsh zone.[18] Additionally, in the same marshes, the reed Phragmites australis has been invading the area expanding to lower marshes and becoming a dominant species. P. australis is an aggressive halophyte that can invade disturbed areas in large numbers outcompeting native plants.[7][36][37] 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.[38][10] The sea level rise causes more open water zones within the salt marsh. These zones cause erosion along their edges, further eroding the marsh into open water until the whole marsh disintegrates.[39]
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.[40] Marshes are susceptible to both erosion and accretion, which play a role in a what is called a bio-geomorphic feedback.[41] 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.[40] However, this feedback is also dependent on other factors like productivity of the vegetation, sediment supply, land subsidence, biomass accumulation, and magnitude and frequency of storms.[40] In a study published by Ü. S. N. Best in 2018,[40] they found that bioaccumulation was the number one factor in a salt marsh's ability to keep up with SLR rates. The salt marsh's resilience depends upon its increase in bed level rate being greater than that of sea levels' increasing rate, otherwise the marsh will be overtaken and drowned.
Biomass accumulation can be measured in the form of above-ground organic biomass accumulation, and below-ground inorganic accumulation by means of sediment trapping and sediment settling from suspension.[42] 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.[42] Along with the vertical accretion of sediment and biomass, the accommodation space for marsh land growth must also be considered. Accommodation space is the land available for additional sediments to accumulate and marsh vegetation to colonize laterally.[43] 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,[44] 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.[45]
Crab herbivory and bioturbation
Crabs, such as the tunnelling mud crab Helice crassa 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.[46]
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.[47] 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.[48] 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.[49]
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.[50] 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.[51] 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, due to not depending upon external organic carbon sources for their growth and survival.[52][53] 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.[54]
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.[55] The dominant class of sulfate-reducing bacteria in salt marshes tends to be Deltaproteobacteria.[55] Some examples of deltaproteobacteria that are found in salt marshes are species of genera Desulfobulbus, Desulfuromonas, and Desulfovibrio.[55]
The abundance and diversity of chemolithoautotrophs in salt marshes is largely determined by the composition of plant species in the salt marsh ecosystem.[55][56] 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,[55] suggested that both the species richness and total abundance of sulfate-reducing bacterial communities increased when a new plant, S. alterniflora, with a higher C-input to the ecosystem was introduced.[55] Although chemolithotrophs produce their own carbon, they still depend on the C-input from salt marshes because of the indirect impact it has on the amount of viable electron donors, such as reduced sulfur compounds.[57] 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.[58]
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.[55]
Why are sulfate-reducing bacteria in salt marshes?
Salt marshes are the ideal environment for sulfate-reducing bacteria. The sulfate-reducing bacteria tend to live in anoxic conditions, such as in salt marshes, because they require reduced compounds to produce their energy.[58] Since there is a high sedimentation rate and a high amount of organic matter, the conditions of the sediment are usually dependably anoxic. However, the conditions all across the salt marsh (above the sediment) are not completely anoxic, which means the organisms living here must have some level of tolerance to oxygen. Many of the chemolithoautotrophs living outside or at the surface of the sediment also exhibit this characteristic.[58]
Significance of sulfate-reducing bacteria
Sulfate-reducing bacteria play a significant role in nutrient recycling and in reducing nitrate pollution levels. Since humans have been adding disproportionate amounts of nitrates to coastal waters, salt marshes are one of the ecosystems where nitrate pollution remains an issue.[59] The enrichment of nitrates in the water increases denitrification, as well as microbial decomposition and primary productivity.[59] Sulfate-reducing and oxidizing bacteria, however, play a role in removing the excess nitrates from the water to prevent eutrophication.[59] Since the sulfate-reducing bacteria is in the water and sediment, reduced sulfur molecules are usually in abundance. These reduced sulfates then react with excess nitrate in the water, reducing nitrate and oxidizing the reduced sulfur.[59] As a result of human nitrate enrichment, it is predicted that sulfur-oxidizing bacteria which also reduce nitrates will increase in relative abundance to sulfur-reducing bacteria.[59]
Abundance and significance of chemolithoautotroph nitrifiers within salt marshes
Within salt marshes, chemolithoautotrophic nitrifying bacteria are also frequently identified, including Betaproteobacteria ammonia oxidizers such as Nitrosomonas and Nitrosospira.[54] Although ammonia-oxidizing Archaea (AOA) are found to be more prevalent than ammonium-oxidizing Bacteria (AOB) within salt marsh environments, predominantly from the Crenarchaeota group, AOB play a critical role within the salt marsh environment too.[60][61] Increases in marsh salinity tend to favor AOB, while higher oxygen levels and lower carbon-to-nitrogen ratios favor AOA.[62] 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-).[63] 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.[64]
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.[54] In addition, the abundance of fixed-nitrogen in these environments critically influences the distribution of the betaproteobacteria within the salt marsh: Nitrosomonas are more found to be in greater abundance within high N and C environments, whereas Nitrosospira are found to be more abundant in lower N and C regions.[54] Further, factors such as temperature, pH, net primary productivity, and regions of anoxia may limit nitrification, and thus critically influence nitrifier distribution.[65]
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.[66] Further, nitrogen oxidation is important for the downstream removal of nitrates into nitrogen gas, catalyzed by denitrifiers, from the marsh environment.[66] Hence, AOB play an indirect role in nitrogen removal into the atmosphere.[66]
Cyanobacteria are important nitrogen fixers in salt marshes, and provide nitrogen to organisms like diatoms and microalgae.[68]
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.[69][70] 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.[71][72] 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.[69] Purple bacteria can be further classified as either purple sulphur bacteria, or purple non-sulfur bacteria. Purple sulphur bacteria are more tolerant to sulfide and store the sulfur they create intracellularly, while purple non-sulfur bacteria excrete any sulfur they produce.
Green bacteria
Green sulfur bacteria (Chlorobiaceae) are photoautotrophic bacteria that utilize sulfide and thiosulfate for their growth, producing sulfate in the process.[73] 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.[74]
Microbial decomposition activity within salt marshes
Phaeosphaeria 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.[78] The carbon from Spartinaalterniflora is made accessible to the salt marsh food web largely through these bacterial communities which are then consumed by bacterivores.[79] Bacteria are responsible for the degradation of up to 88% of lignocellulotic material in salt marshes.[79] However, fungal populations have been found to dominate over bacterial populations in winter months.[80]
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.[78] In terms of bacteria, the alphaproteobacteria class is the most prevalent class within the salt marsh environment involved in decomposition activity.[78] The propagation of Phaeosphaeria spartinicola is through ascospores that are released when the host plant is wetted by high tides or rain.[81]
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.[27] They are now protected by legislation in many countries to prevent the loss of these ecologically important habitats.[82] 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.[7]
Bakker et al. (1997)[83] 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)[83] is to restore the destroyed habitat into its natural state either at the original site or as a replacement at a different site. Under natural conditions, recovery can take 2–10 years or even longer depending on the nature and degree of the disturbance and the relative maturity of the marsh involved.[82] Marshes in their pioneer stages of development will recover more rapidly than mature marshes[82] as they are often first to colonize the land. It is important to note that restoration can often be sped up through the replanting of native vegetation.
Common reed (Phragmites australis) an invasive species in degraded marshes in the northeastern United States.
This last approach is often the most practiced and generally more successful than allowing the area to naturally recover on its own. The salt marshes in the state of Connecticut in the United States have long been an area lost to fill and dredging. As of 1969, the Tidal Wetland Act was introduced that ceased this practice,[37] but despite the introduction of the act, the system was still degrading due to alterations in tidal flow. One area in Connecticut is the marshes on Barn Island. These marshes were diked then impounded with salt and brackish marsh during 1946–1966.[37] As a result, the marsh shifted to a freshwater state and became dominated by the invasive species P. australis, Typha angustifolia and T. latifolia that have little ecological connection to the area.[37]
By 1980, a restoration programme was put in place that has now been running for over 20 years.[37] This programme has aimed to reconnect the marshes by returning tidal flow along with the ecological functions and characteristics of the marshes back to their original state. In the case of Barn Island, reduction of the invasive species has been initiated, re-establishing the tidal-marsh vegetation along with animal species such as fish and insects. This example highlights that considerable time and effort is needed to effectively restore salt marsh systems. The timescale for salt marsh recovery is dependent on the development stage of the marsh, type and extent of the disturbance, geographical location and the environmental and physiological stress factors to the marsh-associated flora and fauna.
Although much effort has gone into restoring salt marshes worldwide, further research is needed. There are many setbacks and problems associated with marsh restoration that require careful long-term monitoring. Information on all components of the salt marsh ecosystem should be understood and monitored from sedimentation, nutrient, and tidal influences, to behaviour patterns and tolerances of both flora and fauna species.[82] Once a better understanding of these processes is acquired, and not just locally, but over a global scale, then more sound and practical management and restoration efforts can be implemented to preserve these valuable marshes and restore them to their original state.
While humans are situated along coastlines, there will always be the possibility of human-induced disturbances despite the number of restoration efforts we plan to implement. Dredging, pipelines for offshore petroleum resources, highway construction, accidental toxic spills or just plain carelessness are examples that will for some time now and into the future be the major influences of salt marsh degradation.[82]
In addition to restoring and managing salt marsh systems based on scientific principles, the opportunity should be taken to educate public audiences of their importance biologically and their purpose as serving as a natural buffer for flood protection.[27] Because salt marshes are often located next to urban areas, they are likely to receive more visitors than remote wetlands. By physically seeing the marsh, people are more likely to take notice and be more aware of the environment around them. An example of public involvement occurred at the Famosa Slough State Marine Conservation Area in San Diego, where a "friends" group worked for over a decade in trying to prevent the area from being developed.[84] Eventually, the 5-hectare (12-acre) 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.[84]
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.[22][23]
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.[25] 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.[23]
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.[20] Marsh surface elevations may be measured with a stadia rod and transit,[23] electronic theodolite,[22] Real-Time Kinematic Global Positioning System,[20]laser level[25] or electronic distance meter (total station). Hydrological dynamics include water depth, measured automatically with a pressure transducer,[22][23][25] or with a marked wooden stake,[21] and water velocity, often using electromagnetic current meters.[21][23]
Eutrophication is a general term describing a process in which nutrients accumulate in a body of water, resulting in an increased growth of microorganisms that may deplete the water of oxygen. Although eutrophication is a natural process, manmade or cultural eutrophication is far more common and is a rapid process caused by a variety of polluting inputs including poorly treated sewage, industrial wastewater, and fertilizer runoff. Such nutrient pollution usually causes algal blooms and bacterial growth, resulting in the depletion of dissolved oxygen in water and causing substantial environmental degradation.
A mangrove is a shrub or tree that grows mainly in coastal saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have special adaptations to take in extra oxygen and to remove salt, which allow them to tolerate conditions that would kill most plants. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in several plant families. They occur worldwide in the tropics and subtropics and even some temperate coastal areas, mainly between latitudes 30° N and 30° S, with the greatest mangrove area within 5° of the equator. Mangrove plant families first appeared during the Late Cretaceous to Paleocene epochs, and became widely distributed in part due to the movement of tectonic plates. The oldest known fossils of mangrove palm date to 75 million years ago.
An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environments and are an example of an ecotone. Estuaries are subject both to marine influences such as tides, waves, and the influx of saline water, and to fluvial influences such as flows of freshwater and sediment. The mixing of seawater and freshwater provides high levels of nutrients both in the water column and in sediment, making estuaries among the most productive natural habitats in the world.
A wetland is a distinct semi-aquatic ecosystem whose groundcovers are flooded or saturated in water, either permanently, for years or decades, or only seasonally for a shorter periods. Flooding results in oxygen-poor (anoxic) processes taking place, especially in the soils. Wetlands form a transitional zone between waterbodies and dry lands, and are different from other terrestrial or aquatic ecosystems due to their vegetation's roots having adapted to oxygen-poor waterlogged soils. They are considered among the most biologically diverse of all ecosystems, serving as habitats to a wide range of aquatic and semi-aquatic plants and animals, with often improved water quality by the plants removing excess nutrients such as nitrates and phosphates.
In ecology, a marsh is a wetland that is dominated by herbaceous plants rather than by woody plants. More in general, the word can be used for any low-lying and seasonally waterlogged terrain. In Europe and in agricultural literature low-lying meadows that require draining and embanked polderlands are also referred to as marshes or marshland.
Coos Bay is an estuary where the Coos River enters the Pacific Ocean, the estuary is approximately 12 miles long and up to two miles wide. It is the largest estuary completely within Oregon state lines. The Coos Bay watershed covers an area of about 600 square miles and is located in northern Coos County, Oregon, in the United States. The Coos River, which begins in the Oregon Coast Range, enters the bay from the east. From Coos River, the bay forms a sharp loop northward before arching back to the south and out to the Pacific Ocean. Haynes Inlet enters the top of this loop. South Slough branches off from the bay directly before its entrance into the Pacific Ocean. The bay was formed when sea levels rose over 20,000 years ago at the end of the Last Glacial Maximum, flooding the mouth of the Coos River. Coos Bay is Oregon's most important coastal industrial center and international shipping port, with close ties to San Francisco, the Columbia River, Puget Sound and other major ports of the Pacific rim.
A tidal creek or tidal channel is a narrow inlet or estuary that is affected by the ebb and flow of ocean tides. Thus, it has variable salinity and electrical conductivity over the tidal cycle, and flushes salts from inland soils. Tidal creeks are characterized by slow water velocity, resulting in buildup of fine, organic sediment in wetlands. Creeks may often be a dry to muddy channel with little or no flow at low tide, but with significant depth of water at high tide. Due to the temporal variability of water quality parameters within the tidally influenced zone, there are unique biota associated with tidal creeks which are often specialised to such zones. Nutrients and organic matter are delivered downstream to habitats normally lacking these, while the creeks also provide access to inland habitat for salt-water organisms.
A tidal marsh is a marsh found along rivers, coasts and estuaries which floods and drains by the tidal movement of the adjacent estuary, sea or ocean. Tidal marshes experience many overlapping persistent cycles, including diurnal and semi-diurnal tides, day-night temperature fluctuations, spring-neap tides, seasonal vegetation growth and decay, upland runoff, decadal climate variations, and centennial to millennial trends in sea level and climate.
An aquatic ecosystem is an ecosystem found in and around a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms—aquatic life—that are dependent on each other and on their environment. The two main types of aquatic ecosystems are marine ecosystems and freshwater ecosystems. Freshwater ecosystems may be lentic ; lotic ; and wetlands.
Mangrove forests, also called mangrove swamps, mangrove thickets or mangals, are productive wetlands that occur in coastal intertidal zones. Mangrove forests grow mainly at tropical and subtropical latitudes because mangroves cannot withstand freezing temperatures. There are about 80 different species of mangroves, all of which grow in areas with low-oxygen soil, where slow-moving waters allow fine sediments to accumulate.
Sporobolus alterniflorus, or synonymously known as Spartina alterniflora, the smooth cordgrass, saltmarsh cordgrass, or salt-water cordgrass, is a perennial deciduous grass which is found in intertidal wetlands, especially estuarine salt marshes. It has been reclassified as Sporobolus alterniflorus after a taxonomic revision in 2014, but it is still common to see Spartina alterniflora and in 2019 an interdisciplinary team of experts coauthored a report published in the journal Ecology supporting Spartina as a genus. It grows 1–1.5 m tall and has smooth, hollow stems that bear leaves up to 20–60 cm long and 1.5 cm wide at their base, which are sharply tapered and bend down at their tips. Like its relative saltmeadow cordgrass S. patens, it produces flowers and seeds on only one side of the stalk. The flowers are a yellowish-green, turning brown by the winter. It has rhizoidal roots, which, when broken off, can result in vegetative asexual growth. The roots are an important food resource for snow geese. It can grow in low marsh as well as high marsh, but it is usually restricted to low marsh because it is outcompeted by salt meadow cordgrass in the high marsh. It grows in a wide range of salinities, from about 5 psu to marine, and has been described as the "single most important marsh plant species in the estuary" of Chesapeake Bay. It is described as intolerant of shade.
Judith Shulman Weis is an American marine biologist. Her research and writing focuses on estuarine ecology and ecotoxicology, including the responses of salt marsh and brackish marsh organisms, populations and communities to stresses, particularly heavy metal contaminants, invasive species and parasites. She is also working to reduce the spread of microplastics in the environment and find solutions to protecting coastal marshes from sea level rise.
High salt marsh dieback, or salt marsh browning, is the primary force in salt marsh degradation in the high marsh. The general effect is that the plants in the marsh die off and brown, leaving dead organic matter, and ultimately open sediment. Without strong plant roots holding the sediment, these open areas of land erode, causing the salt marsh to retreat back to the mainland. Dieback zones lack their main producers, such as the salt marsh cord grass, or Spartina alterniflora, and ultimately become completely unproductive.
The Coastal and Estuarine Research Federation (CERF) is a private, nonprofit organization created in 1971. At that time, the members of two regionally based organizations, the Atlantic Estuarine Research Society (AERS) and the New England Estuarine Research Society (NEERS) recognized the need for a third estuarine organization that would address national estuarine and coastal issues. Today, CERF is a multidisciplinary federation of members and seven regionally-based Affiliate Societies dedicated to the understanding and wise stewardship of estuaries and coasts worldwide.
Brackish marshes develop from salt marshes where a significant freshwater influx dilutes the seawater to brackish levels of salinity. This commonly happens upstream from salt marshes by estuaries of coastal rivers or near the mouths of coastal rivers with heavy freshwater discharges in the conditions of low tidal ranges.
Low marsh is a tidal marsh zone located below the Mean Highwater Mark (MHM). Based on elevation, frequency of submersion, soil characteristics, vegetation, microbial community, and other metrics, salt marshes can be divided to into three distinct areas: low marsh, middle marsh/high marsh, and the upland zone. Low marsh is characterized as being flooded daily with each high tide, while remaining exposed during low tides.
Geukensia demissa is a species of mussel, a marine bivalve mollusk in the family Mytilidae, the true mussels. This species is native to the Atlantic coast of North America. The common names for this species include ribbed mussel, Atlantic ribbed marsh mussel and ribbed horsemussel. However, the common name ribbed mussel is also used for the Southern Hemisphere mussel Aulacomya atra. The appearance of the shell is grooved and oval in shape. The interior of this mussel is tinted purple.
Juncus roemerianus is a species of flowering plant in the rush family known by the common names black rush, needlerush, and black needlerush. It is native to North America, where its main distribution lies along the coastline of the southeastern United States, including the Gulf Coast. It occurs from New Jersey to Texas, with outlying populations in Connecticut, New York, Mexico, and certain Caribbean islands.
Salt marsh die-off is a term that has been used in the US and UK to describe the death of salt marsh cordgrass leading to subsequent degradation of habitat, specifically in the low marsh zones of salt marshes on the coasts of the Western Atlantic. Cordgrass normally anchors sediment in salt marshes; its loss leads to decreased substrate hardness, increased erosion, and collapse of creek banks into the water, ultimately resulting in decreased marsh health and productivity.
The marsh organ is a collection of plastic pipes attached to a wooden framework that is placed in marshes to measure the effects of inundation time and flood frequency on the productivity of marsh vegetation. The information is used for scientific research purposes.
↑ Allen, JRL, Pye, K (1992). Saltmarshes: morphodynamics, conservation, and engineering significance. Cambridge University Press. Cambridge, UK.
1 2 3 Chapman, V. J. (1974). Salt marshes and salt deserts of the world. Phyllis Claire Chapman, Germany.
1 2 3 4 5 6 7 8 Bromberg-Gedan, K., Silliman, B. R., and Bertness, M. D. (2009). "Centuries of human driven change in salt marsh ecosystems", Annual Review of Marine Science, 1: 117–141.
↑ Vernberg, F. J. 1993. Salt-Marsh Processes: A Review. Environmental Toxicology and Chemistry 12:2167–2195.
1 2 Scott, D. B., J. Frail-Gauthier, and P. J. Mudie. 2014. Coastal wetlands of the world: geology, ecology, distribution and applications. Cambridge University Press, New York
1 2 3 4 5 Pethick, J. (1984). An introduction to coastal geomorphology. Edward Arnold, London.
1 2 3 Boorman, L., Hazelden, J., and Boorman, M. (2002). "New salt marshes for old – salt marsh creation and management". The Changing Coast, EUROCAST/EUCC, EUROCOAST Littoral 2002: Porto, Portugal; 35–45.
↑ Ginsburg, R. N., and Lowenstam, H. A. (1958). "The influence of marine bottom communities on the depositional environment of sediments". The Journal of Geology, 66: (3), 310–318.
↑ Aspden, R. J., Vardy, S. and Paterson, D. M. (2004). Salt marsh microbial ecology: microbes, benthic mats and sediment movement. In Fagherazzi, S., Marani, M. and Blum, L. K. (Eds), The Ecogeomorphology of Tidal Marshes (pp. 115–136). American Geophysical Union, Washington, DC.
1 2 3 Bird, E. (2008). Coastal geomorphology: an introduction. John Wiley & Sons Ltd, West Sussex, England.
1 2 3 4 5 Bertness, MD, Ewanchuk, PJ, Silliman, BR (2002). "Anthropogenic modification of New England salt marsh landscapes". Proceedings of the National Academy of Sciences99(3): 1395–1398.
1 2 Rand, TA (2000). Seed Dispersal, Habitat Suitability and the Distribution of Halophytes across a Salt Marsh Tidal Gradient. Journal of Ecology88(4): 608–621.
1 2 3 4 5 6 Li, H. and Yang, S. L. (2000). "Trapping effect of tidal marsh vegetation on suspended sediment, Yangtze Delta". Journal of Coastal Research, 25: (4), 915–924
1 2 3 4 5 6 Shi, Z., Hamilton, L. J. and Wolanski, E. (2000). "Near-bed currents and suspended sediment transport in saltmarsh canopies". Journal of Coastal Research, 16: (3), 908–914.
1 2 3 4 5 6 7 8 Reed, D. J., Spencer, T., Murray, A. L., French, J. R. and Leonard, L. (1999). "Marsh surface sediment deposition and the role of tidal creeks: implications for created and managed coastal marshes". Journal of Coastal Conservation, 5: (1), 81–90.
1 2 3 4 5 6 7 Wood, N. and Hine, A. C. (2007). "Spatial trends in marsh sediment deposition within a microtidal creek system, Wacasassa Bay, Florida". Journal of Coastal Research, 23: (4), 823–833.
1 2 3 4 Cahoon, D. R., White, D. A. and Lynch, J. C. (2011). "Sediment infilling and wetland formation dynamics in an active crevasse splay of the Mississippi River delta". Geomorphology, 131: 57–68.
↑ Hinde, HP (1954). "The Vertical Distribution of Salt Marsh Phanerogams in Relation to Tide Levels". Ecological Monographs24(2): 209–225.
1 2 3 King, SE, Lester, JN (1995). "The Value of Salt Marsh as a Sea Defence". Marine Pollution Bulletin30(3): 180–189.
1 2 3 Long, S. P. and Mason, C. F. (1983). Saltmarsh ecology. Blackie & Son Ltd, Glasgow.
↑ French, J. R. and Burningham, H. (2003). "Tidal marsh sedimentation versus sea-level rise: a southeast England estuarine perspective", Proceedings Coastal Sediments, 1–13.
↑ Angus, G. and Wolters, M. (2008). "The natural regeneration of salt marsh on formerly reclaimed land". Applied Vegetation Science, 11: 335–344.
1 2 Ranwell, D. S. (1972). Ecology of salt marshes and sand dunes. Chapman and Hall Ltd, London.
↑ Kirwan, M. L., Murray, A. B., Donnelly, J. P. and Corbett, D. (2011). "Rapid wetland expansion during European settlement and its implication for marsh survival under modern sediment delivery rates". Geological Society of America, 39: (5), 507–510.
↑ Jupp, K. (2007). Establishing a physical and biological basis for salt marsh restoration and management in the Avon-Heathcote Estuary. Christchurch, University of Canterbury.
↑ Langis, R, Zalejko, M, Zedler, JB (1991). "Nitrogen Assessments in a Constructed and a Natural Salt Marsh of San Diego Bay". Ecological Applications1(1): 40–51.
↑ Chambers, RM, Meyerson, LA, Saltonstall, K (1999). "Expansion of Phragmites australis into tidal wetlands of North America". Aquatic Botany64: 261–273.
1 2 3 4 5 Warren, RS, Fell, PE, Rozsa, R, Brawley, AH, Orsted, AC, Olson, ET, Swamy, V, Niering, WA (2002). "Salt Marsh Restoration in Connecticut: 20 years of Science and Management". Restoration Ecology10(3): 497–513.
↑ Alberti, J., Cebrian, J., Casariego, A. M., Canepuccia, A., Escapa, M. and Iribarne, O. (2011). "Effects of nutrient enrichment and crab herbivory on a SW Atlantic salt marsh" productivity. Journal of Experimental Marine Biology and Ecology, 405: 99–104.
↑ Holdredge, C., Bertness, M. D. and Altieri, A. H. (2008). "Role of crab herbivory in die-off of New England salt marshes". Conservation Biology, 23: (3), 672–679.
↑ Smith, S. M. and Tyrrell, M. C. (2012). "Effects of mud fiddler crabs (Uca pugnax) on the recruitment of halophyte seedlings in salt marsh dieback areas of Cape Cod" (Massachusetts, US). Ecological Research, 27: 233–237.
1 2 Bakker, JP, Esselink, P, Van Der Wal, R, Dijkema, KS (1997). 'Options for restoration and management of coastal salt marshes in Europe,' in Urbanska, KM, Webb, NR, Edwards, PJ (eds), Restoration Ecology and Sustainable Development. Cambridge University Press, UK. p. 286-322.
1 2 Callaway, JC, Zedler, JB (2004). "Restoration of urban salt marshes: Lessons from southern California". Urban Ecosystems7: 107–124.
This page is based on this Wikipedia article Text is available under the CC BY-SA 4.0 license; additional terms may apply. Images, videos and audio are available under their respective licenses.
