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A fen is a type of
peat Peat (), also known as turf (), is an accumulation of partially decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, moors, or muskegs. The peatland ecosystem covers and is the most efficien ...
-accumulating
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...
fed by mineral-rich
ground Ground may refer to: Geology * Land, the surface of the Earth not covered by water * Soil, a mixture of clay, sand and organic matter present on the surface of the Earth Electricity * Ground (electricity), the reference point in an electrical c ...
or
surface water Surface water is water located on top of land forming terrestrial (inland) waterbodies, and may also be referred to as ''blue water'', opposed to the seawater and waterbodies like the ocean. The vast majority of surface water is produced by prec ...
. It is one of the main types of wetlands along with
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found at ...
es,
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
s, and
bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg; a ...
s. Bogs and fens, both peat-forming ecosystems, are also known as mires. The unique water chemistry of fens is a result of the ground or surface water input. Typically, this input results in higher mineral concentrations and a more
basic BASIC (Beginners' All-purpose Symbolic Instruction Code) is a family of general-purpose, high-level programming languages designed for ease of use. The original version was created by John G. Kemeny and Thomas E. Kurtz at Dartmouth College ...
pH than found in bogs. As peat accumulates in a fen,
groundwater Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidate ...
input can be reduced or cut off, making the fen ombrotrophic rather than
minerotrophic Minerotrophic refers to environments that receive nutrients primarily through groundwater that flows through mineral-rich soils or rock,Environment Canada (2014). Ontario wetland evaluation system: Northern Manual, 1st edition, version 3.2. Queen’ ...
. In this way, fens can become more acidic and transition to bogs over time. Fens can be found around the world, but the vast majority are located at the mid to high latitudes of the Northern Hemisphere. They are dominated by sedges and
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es, particularly
graminoid In botany and ecology, graminoid refers to a herbaceous plant with a grass-like morphology, i.e. elongated culms with long, blade-like leaves. They are contrasted to forbs, herbaceous plants without grass-like features. The plants most ofte ...
s that may be rarely found elsewhere, such as the sedge species '' Carex exilis''. Fens are highly biodiverse ecosystems and often serve as habitats for endangered or rare species, with species composition changing with water chemistry. They also play important roles in the cycling of nutrients such as carbon, nitrogen, and
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
due to the lack of oxygen (anaerobic conditions) in waterlogged organic fen soils. Fens have historically been converted to agricultural land. However, fens face a number of other threats, including peat cutting, pollution, invasive species, and nearby disturbances that lower the water table in the fen, such as quarrying. Interrupting the flow of mineral-rich water into a fen changes the water chemistry, which can alter species richness and dry out the peat. Drier peat is more easily decomposed and can even burn.


Distribution and extent

Fens are distributed around the world, but are most frequently found at the mid-high latitudes of the Northern Hemisphere. They are found throughout the
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
zone and
boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...
regions, but are also present in
tundra In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless moun ...
and in specific environmental conditions in other regions around the world. In the United States, fens are most common in the Midwest and Northeast, but can be found across the country. In Canada, fens are most frequent in the lowlands near
Hudson Bay Hudson Bay ( crj, text=ᐐᓂᐯᒄ, translit=Wînipekw; crl, text=ᐐᓂᐹᒄ, translit=Wînipâkw; iu, text=ᑲᖏᖅᓱᐊᓗᒃ ᐃᓗᐊ, translit=Kangiqsualuk ilua or iu, text=ᑕᓯᐅᔭᕐᔪᐊᖅ, translit=Tasiujarjuaq; french: b ...
and
James Bay James Bay (french: Baie James; cr, ᐐᓂᐯᒄ, Wînipekw, dirty water) is a large body of water located on the southern end of Hudson Bay in Canada. Both bodies of water extend from the Arctic Ocean, of which James Bay is the southernmost par ...
, but can also be found across the country. Fens are also spread across the northern latitudes of Eurasia, including Britain and Ireland, as well as Japan, but east-central Europe is especially rich in fens. Further south, fens are much rarer, but do exist under specific conditions. In Africa, fens have been found in the Okavango Delta in Botswana and the highland slopes in
Lesotho Lesotho ( ), officially the Kingdom of Lesotho, is a country landlocked country, landlocked as an Enclave and exclave, enclave in South Africa. It is situated in the Maloti Mountains and contains the Thabana Ntlenyana, highest mountains in Sou ...
. Fens can also be found at the colder latitudes of the Southern Hemisphere. They are found in New Zealand and southwest Argentina, but the extent is much less than that of the northern latitudes. Locally, fens are most often found at the intersection of terrestrial and aquatic ecosystems, such as the headwaters of streams and rivers. It is estimated that there are approximately 1.1 million square kilometers of fens worldwide, but quantifying the extent of fens is difficult. Because wetland definitions vary regionally, not all countries define fens the same way. In addition, wetland data is not always available or of high quality. Fens are also difficult to rigidly delineate and measure, as they are located between terrestrial and aquatic ecosystems.


Definition

Rigidly defining types of wetlands, including fens, is difficult for a number of reasons. First, wetlands are diverse and varied ecosystems that are not easily categorized according to inflexible definitions. They are often described as a transition between terrestrial and aquatic ecosystems with characteristics of both. This makes it difficult to delineate the exact extent of a wetland. Second, terms used to describe wetland types vary greatly by region. The term ''
bayou In usage in the Southern United States, a bayou () is a body of water typically found in a flat, low-lying area. It may refer to an extremely slow-moving stream, river (often with a poorly defined shoreline), marshy lake, wetland, or creek. They ...
,'' for example, describes a type of wetland, but its use is generally limited to the southern United States. Third, different languages use different terms to describe types of wetlands. For instance, in Russian, there is no equivalent word for the term ''
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
'' as it is typically used in North America. The result is a large number of
wetland classification Classification of wetlands has been a problematical task, with the commonly accepted definition of what constitutes a wetland being among the major difficulties. A number of national wetland classifications exist. In the 1970s, the Ramsar Conventio ...
systems that each define wetlands and wetland types in their own way. However, many classification systems include four broad categories that most wetlands fall into:
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found at ...
, swamp,
bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg; a ...
, and fen. While classification systems differ on the exact criteria that define a fen, there are common characteristics that describe fens generally and imprecisely. A general definition provided by the textbook ''Wetlands'' describes a fen as "a peat-accumulating wetland that receives some drainage from surrounding mineral soil and usually supports marsh like vegetation." Three examples are presented below to illustrate more specific definitions for the term ''fen''.


Canadian Wetland Classification System definition

In the Canadian Wetland Classification System, fens are defined by six characteristics: # Peat is present. # The surface of the wetland is level with the
water table The water table is the upper surface of the zone of saturation. The zone of saturation is where the pores and fractures of the ground are saturated with water. It can also be simply explained as the depth below which the ground is saturated. T ...
. Water flows on the surface and through the subsurface of the wetland. # The water table fluctuates. It may be at the surface of the wetland or a few centimeters above or below it. # The wetland receives a significant amount of its water from mineral-rich groundwater or surface water. # Decomposed sedges or brown moss peat are present. # The vegetation is predominantly
graminoid In botany and ecology, graminoid refers to a herbaceous plant with a grass-like morphology, i.e. elongated culms with long, blade-like leaves. They are contrasted to forbs, herbaceous plants without grass-like features. The plants most ofte ...
s and shrubs.


''Wetland Ecology: Principles and Conservation'' (Keddy) definition

In the textbook ''Wetland Ecology: Principles and Conservation'', Paul A. Keddy offers a somewhat simpler definition of a fen as "a wetland that is usually dominated by sedges and grasses rooted in shallow peat, often with considerable groundwater movement, and with pH greater than 6." This definition differentiates fens from swamps and marshes by the presence of peat.


''The Biology of Peatlands'' (Rydin) definition

In ''The Biology of Peatlands'' fens are defined by the following criteria: # The wetland is not flooded by lake or stream water. # Woody vegetation 2 meters or taller is absent or canopy cover is less than 25%. # The wetland is
minerotrophic Minerotrophic refers to environments that receive nutrients primarily through groundwater that flows through mineral-rich soils or rock,Environment Canada (2014). Ontario wetland evaluation system: Northern Manual, 1st edition, version 3.2. Queen’ ...
(it receives its nutrients from mineral-rich groundwater). A further distinction is made between open and wooded fens, where open fens have canopy cover less than 10% and wooded fens have 10–25% canopy cover. If tall shrubs or trees dominate, the wetland is instead classified as a wooded bog or
swamp forest Freshwater swamp forests, or flooded forests, are forests which are inundated with freshwater, either permanently or seasonally. They normally occur along the lower reaches of rivers and around freshwater lakes. Freshwater swamp forests are found ...
, depending on other criteria.


Biogeochemical features


Hydrological conditions

Hydrology Hydrology () is the scientific study of the movement, distribution, and management of water on Earth and other planets, including the water cycle, water resources, and environmental watershed sustainability. A practitioner of hydrology is calle ...
al conditions, as seen in other wetlands, are a major determinant of fen biota and
biogeochemistry Biogeochemistry is the scientific discipline that involves the study of the chemical, physical, geological, and biological processes and reactions that govern the composition of the natural environment (including the biosphere, the cryosphere, t ...
. Fen soils are constantly inundated because the water table is at or near the surface. The result is anaerobic (oxygen-free) soils due to the slow rate at which oxygen diffuses into waterlogged soil. Anaerobic soils are ecologically unique because earth's atmosphere is oxygenated, while most terrestrial ecosystems and surface waters are aerobic. The anaerobic conditions found in wetland soils result in reduced, rather than
oxidized Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a d ...
, soil chemistry. A hallmark of fens is that a significant portion of their water supply is derived from
groundwater Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidate ...
(minerotrophy). Because hydrology is the dominant factor in wetlands, the chemistry of the groundwater has an enormous effect on the characteristics of the fen it supplies. Groundwater chemistry, in turn, is largely determined by the geology of the rocks that the groundwater flows through. Thus, the characteristics of a fen, especially its pH, are directly influenced by the type of rocks its groundwater supply contacts. pH is a major factor in determining fen species composition and richness, with more basic fens called "rich" and more acidic fens called "poor." Rich fens tend to be highly biodiverse and harbor a number of rare or endangered species, and biodiversity tends to decrease as the richness of fen decreases. Fens tend to be found above rocks that are rich in calcium, such as
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
. When groundwater flows past calcareous (calcium-rich) rocks like limestone ( calcium carbonate), a small amount dissolves and is carried to the fen supplied by the groundwater. When calcium carbonate dissolves, it produces
bicarbonate In inorganic chemistry, bicarbonate (IUPAC-recommended nomenclature: hydrogencarbonate) is an intermediate form in the deprotonation of carbonic acid. It is a polyatomic anion with the chemical formula . Bicarbonate serves a crucial biochemic ...
and a
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to ...
cation An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by convent ...
according to the following equilibrium: CaCO3 + H2CO3 <=> Ca^2+ + 2HCO3^- where carbonic acid (H2CO3) is produced by the dissolution of
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
in water. In fens, the bicarbonate anion produced in this equilibrium acts as a pH buffer, which keeps the pH of the fen relatively stable. Fens supplied by groundwater that doesn't flow through minerals and act as a
buffer Buffer may refer to: Science * Buffer gas, an inert or nonflammable gas * Buffer solution, a solution used to prevent changes in pH * Buffering agent, the weak acid or base in a buffer solution * Lysis buffer, in cell biology * Metal ion buffer * ...
when dissolved tend to be more acidic. The same effect is observed when groundwater flows through minerals with low solubility, such as sand. In extreme rich fens, calcium carbonate can precipitate out of solution to form
marl Marl is an earthy material rich in carbonate minerals, clays, and silt. When hardened into rock, this becomes marlstone. It is formed in marine or freshwater environments, often through the activities of algae. Marl makes up the lower part o ...
deposits. Calcium carbonate precipitates out of solution when the
partial pressure In a mixture of gases, each constituent gas has a partial pressure which is the notional pressure of that constituent gas as if it alone occupied the entire volume of the original mixture at the same temperature. The total pressure of an ideal gas ...
of carbon dioxide in the solution falls. The decrease in carbon dioxide partial pressure is caused by uptake by plants for photosynthesis or direct loss to the atmosphere. This reduces the availability of carbonic acid in solution, shifting the above equilibrium back towards the formation of calcium carbonate. The result is the precipitation of calcium carbonate and the formation of marl.


Nutrient cycling

Fen, being a distinct type of wetland, shares many biogeochemical characteristics with other wetlands. Like all wetlands, they play an important role in
nutrient cycling A nutrient cycle (or ecological recycling) is the movement and exchange of inorganic and organic matter back into the production of matter. Energy flow is a unidirectional and noncyclic pathway, whereas the movement of mineral nutrients is cycli ...
because they are located at the interface of aerobic (oxic) and anaerobic (anoxic) environments. Most wetlands have a thin top layer of oxygenated soil in contact with the atmosphere or oxygenated surface waters. Nutrients and minerals may cycle between this oxidized top layer and the reduced layer below, undergoing oxidation and reduction reactions by the microbial communities adapted to each layer. Many important reactions take place in the reduced layer, including
denitrification Denitrification is a microbially facilitated process where nitrate (NO3−) is reduced and ultimately produces molecular nitrogen (N2) through a series of intermediate gaseous nitrogen oxide products. Facultative anaerobic bacteria perform denitr ...
, manganese reduction, iron reduction, sulfate reduction, and
methanogenesis Methanogenesis or biomethanation is the formation of methane coupled to energy conservation by microbes known as methanogens. Organisms capable of producing methane for energy conservation have been identified only from the domain Archaea, a group ...
. Because wetlands are hotspots for nutrient transformations and often serve as nutrient sinks, they may be constructed to treat nutrient-rich waters created by human activities. Fens are also hotspots for
primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through c ...
, as the continuous input of groundwater stimulates production.
Bog A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and muskeg; a ...
s, which lack this input of
groundwater Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidate ...
, have much lower primary production.


Carbon

Carbon from all types of wetlands, including fens, arrives mostly as
organic carbon Total organic carbon (TOC) is the amount of carbon found in an organic compound and is often used as a non-specific indicator of water quality or cleanliness of pharmaceutical manufacturing equipment. TOC may also refer to the amount of organic c ...
from either adjacent upland ecosystems or by photosynthesis in the wetland itself. Once in the wetland, organic carbon generally has three main fates: oxidation to CO2 by
aerobic respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
, burial as organic matter in peat, or decomposition to
methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The relative abundance of methane on Eart ...
. In peatlands, including fens, primary production by plants is greater than decomposition, which results in the accumulation of organic matter as peat. Resident mosses usually carry out decomposition within the fen, and temperate fens are often driven by plant roots' decomposition. These peat stores sequester an enormous amount of carbon. Nevertheless, it is difficult to determine whether fens net take up or emit
greenhouse gas A greenhouse gas (GHG or GhG) is a gas that Absorption (electromagnetic radiation), absorbs and Emission (electromagnetic radiation), emits radiant energy within the thermal infrared range, causing the greenhouse effect. The primary greenhouse ...
es. This is because fens emit methane, which is a more potent greenhouse gas than carbon dioxide. Methanogenic
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
that reside in the anaerobic layers of peat combine carbon dioxide and hydrogen gas to form methane and water. This methane can then escape into the atmosphere and exert its warming effects. Peatlands dominated by brown mosses and sedges such as fens have been found to emit a greater amount of methane than '' Sphagnum''-dominated peatlands such as bogs.


Nitrogen

Fens play an important role in the global
nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmospheric, terrestrial, and marine ecosystems. The conversion of nitrogen can be carried out through both biologi ...
due to the anaerobic conditions found in their soils, which facilitate the oxidation or reduction of one form of nitrogen to another. Most nitrogen arrives in wetlands as nitrate from
runoff Runoff, run-off or RUNOFF may refer to: * RUNOFF, the first computer text-formatting program * Runoff or run-off, another name for bleed, printing that lies beyond the edges to which a printed sheet is trimmed * Runoff or run-off, a stock market ...
, in organic matter from other areas, or by
nitrogen fixation Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmo ...
in the wetland. There are three main forms of nitrogen found in wetlands: nitrogen in organic matter, oxidized nitrogen (
nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...
or
nitrite The nitrite polyatomic ion, ion has the chemical formula . Nitrite (mostly sodium nitrite) is widely used throughout chemical and pharmaceutical industries. The nitrite anion is a pervasive intermediate in the nitrogen cycle in nature. The name ...
), and
ammonium The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternary a ...
. Nitrogen is abundant in peat. When the organic matter in peat is decomposed in the absence of oxygen, ammonium is produced via
ammonification The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmospheric, terrestrial, and marine ecosystems. The conversion of nitrogen can be carried out through both biolo ...
. In the oxidized surface layer of the wetland, this ammonium is oxidized to nitrite and nitrate by
nitrification ''Nitrification'' is the biological oxidation of ammonia to nitrite followed by the oxidation of the nitrite to nitrate occurring through separate organisms or direct ammonia oxidation to nitrate in comammox bacteria. The transformation of amm ...
. The production of ammonium in the reduced layer and its consumption in the top oxidized layer drives upward
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
of ammonium. Likewise, nitrate production in the oxidized layer and nitrate consumption in the reduced layer by denitrification drives downward diffusion of nitrate.
Denitrification Denitrification is a microbially facilitated process where nitrate (NO3−) is reduced and ultimately produces molecular nitrogen (N2) through a series of intermediate gaseous nitrogen oxide products. Facultative anaerobic bacteria perform denitr ...
in the reduced layer produces nitrogen gas and some
nitrous oxide Nitrous oxide (dinitrogen oxide or dinitrogen monoxide), commonly known as laughing gas, nitrous, or nos, is a chemical compound, an oxide of nitrogen with the formula . At room temperature, it is a colourless non-flammable gas, and has a ...
, which then exit the wetland to the atmosphere. Nitrous oxide is a potent greenhouse gas whose production is limited by nitrate and nitrite concentrations in fens. Nitrogen, along with phosphorus, controls how fertile a wetland is.


Phosphorus

Almost all of the phosphorus that arrives in a wetland does so through sediments or plant litter from other ecosystems. Along with nitrogen, phosphorus limits wetland fertility. Under basic conditions like those found in extremely rich fens, calcium will bind to
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phospho ...
anions to make calcium phosphates, which are unavailable for uptake by plants. Mosses also play a considerable role in aiding plants in phosphorus uptake by decreasing soil phosphorus stress and stimulating
phosphatase In biochemistry, a phosphatase is an enzyme that uses water to cleave a phosphoric acid Ester, monoester into a phosphate ion and an Alcohol (chemistry), alcohol. Because a phosphatase enzyme catalysis, catalyzes the hydrolysis of its Substrate ...
activity in organisms found below the moss cover. Helophytes have been shown to bolster phosphorus cycling within fens, especially in fen reestablishment, due to their ability to act as a phosphorus sink, which prevents residual phosphorus in the fen from being transferred away from the it. Under normal conditions, phosphorus is held within soil as dissolved inorganic phosphorus, or
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phospho ...
, which leaves trace amounts of phosphorus in the rest of the ecosystem. Iron is important in phosphorus cycling within fens. Iron can bind to high levels of inorganic phosphate within the fen, leading to a toxic environment and inhibition of plant growth. In iron-rich fens, the area can become vulnerable to acidification, excess nitrogen and potassium, and low water levels. Peat soils play a role in preventing the bonding of irons to phosphate by providing high levels of organic anions for iron to bind to instead of inorganic anions such as phosphate.


Bog-rich fen gradient

Bogs and fens can be thought of as two ecosystems on a gradient from poor to rich, with bogs at the poor end, extremely rich fens at the rich end, and poor fens in between. In this context, "rich" and "poor" refer to the species richness, or how biodiverse a fen or bog is. The richness of these species is strongly influenced by pH and concentrations of calcium and bicarbonate. These factors assist in identifying where along the gradient a particular fen falls. In general, rich fens are
minerotrophic Minerotrophic refers to environments that receive nutrients primarily through groundwater that flows through mineral-rich soils or rock,Environment Canada (2014). Ontario wetland evaluation system: Northern Manual, 1st edition, version 3.2. Queen’ ...
, or dependent on mineral-rich groundwater, while bogs are ombrotrophic, or dependent on precipitation for water and nutrients. Poor fens fall between these two.


Rich fens

Rich fens are strongly minerotrophic; that is, a large proportion of their water comes from mineral-rich ground or surface water. Fens that are more distant from surface waters such as rivers and lakes, however, have been shown to be more rich than fens that are connected. This water is dominated by calcium and bicarbonate, resulting in a slightly acidic to slightly basic pH, which is characteristic of rich fens. These conditions promote high biodiversity. Within rich fens, there is a large amount of variability. The richest fens are the extreme rich (marl) fens, where marl deposits are often build up. These are often pH 7 or greater. Rich and intermediate rich fens are generally neutral to slightly acidic, with a pH of approximately 7 to 5. Rich fens are not always very productive; at high calcium concentrations, calcium ions bind to phosphate anions, reducing the availability of phosphorus and decreasing primary production. Rich bogs with limited primary production can stabilize with the accumulation of mosses and
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
, which promote phosphorus cycling and can support the growth of new vegetation and bacteria. Brown mosses (family '' Amblystegiaceae'') and sedges (genus '' Carex'') are the dominant vegetation. However, an accumulation of mosses such as '' Sphagnum'' can lead to the acidification of the rich fen, potentially converting it into a poor fen. Compared to poor fens, rich fens have higher concentrations of bicarbonate, base cations (Na+, Ca2+, K+, Mg2+), and sulfate.


Poor fens

Poor fens are in many ways an intermediate between rich fens and bogs. Hydrologically, they are more alike to rich fens than to bogs, but in terms of vegetation composition and chemistry, they are more similar to bogs than rich fens. They are much more acidic than their rich counterparts, with a pH of approximately 5.5 to 4. Peat in poor fens tends to be thicker than that of rich fens, which cuts off vegetation access to the mineral-rich soil underneath. In addition, the thicker peat reduces the influence of mineral-rich groundwater that buffers the pH. This makes the fen more ombrotrophic, or dependent on nutrient-poor precipitation for its water and nutrients. Poor fens may also form in areas where the groundwater supplying the fen flows through sediments that don't dissolve well or have low buffering capacity when dissolved. Species richness tends to be lower than that of rich fens but higher than that of bogs. Poor fens, like bogs, are dominated by ''Sphagnum'' mosses, which acidify the fen and decrease nutrient availability.


Threats

One of the many threats that fens face is conversion to agricultural lands. Where climates are suitable, fens have been drained for agricultural use alongside crop production,
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to roam around and consume wild vegetations in order to convert the otherwise indigestible (by human gut) cellulose within grass and other ...
, and hay making. Draining a fen directly is particularly damaging because it lowers the water table. A lower water table can increase aeration and dry out peat, allowing for aerobic decomposition or burning of the organic matter in peat. Draining a fen indirectly by decreasing its water supply can be just as damaging. Disrupting groundwater flow into the fen with nearby human activities such as quarrying or residential development changes how much water and nutrients enter the fen. This can make the fen more ombrotrophic (dependent on precipitation), which results in acidification and a change in water chemistry. This has a direct impact on the habitat of these species and many signature fen species disappear. Fens are also threatened by
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
,
fragmentation Fragmentation or fragmented may refer to: Computers * Fragmentation (computing), a phenomenon of computer storage * File system fragmentation, the tendency of a file system to lay out the contents of files non-continuously * Fragmented distributi ...
, peat cutting, and pollution. Non-native invasive species, such as the
common buckthorn ''Rhamnus cathartica'', the European buckthorn, common buckthorn, purging buckthorn, or just buckthorn, is a species of small tree in the flowering plant family Rhamnaceae. It is native to Europe, northwest Africa and western Asia, from the cent ...
in North America, can invade fens and outcompete rare fen species, reducing biodiversity.
Habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological processes ...
threatens fen species, especially rare or endangered species that are unable to move to nearby fens due to fragmentation. Peat cutting, while much more common in bogs, does happen in fens. Peat cut from fens has many uses, including burning as a fuel. Pollutants can alter the chemistry of fens and facilitate invasion by
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
. Common pollutants of fens include road salts, nutrients from
septic tank A septic tank is an underground chamber made of concrete, fiberglass, or plastic through which domestic wastewater (sewage) flows for basic sewage treatment. Settling and anaerobic digestion processes reduce solids and organics, but the treatme ...
s, and runoff of agricultural fertilizers and pesticides.


Use of term in literature

Shakespeare used the term "fen-sucked" to describe the fog (literally: rising from marshes) in ''
King Lear ''King Lear'' is a tragedy written by William Shakespeare. It is based on the mythological Leir of Britain. King Lear, in preparation for his old age, divides his power and land between two of his daughters. He becomes destitute and insane an ...
'', when Lear says "Infect her beauty, You fen-sucked fogs drawn by the powerful sun, To fall and blister."


Images

File:Kakerdaja fen in spring.JPG, Kakerdaja Fen, Estonia File:Dernford Fen 4.jpg,
Dernford Fen Dernford Fen is a biological Site of Special Scientific Interest north-west of Sawston in Cambridgeshire. The site is a rare surviving example of rough fen A fen is a type of peat-accumulating wetland fed by mineral-rich ground or surface ...
, Cambridgeshire File:Sugar Fen 7.jpg, Sugar Fen, Norfolk File:Illustrated diagram of a fen.jpg


See also

*
Appalachian bogs Appalachian bogs are boreal or hemiboreal ecosystems, which occur in many places in the Appalachian Mountains, particularly the Allegheny and Blue Ridge subranges. Though popularly called bogs, many of them are technically fens. Natural histo ...
* List of fen plants *
Bayou In usage in the Southern United States, a bayou () is a body of water typically found in a flat, low-lying area. It may refer to an extremely slow-moving stream, river (often with a poorly defined shoreline), marshy lake, wetland, or creek. They ...
* Biodiversity Action Plan * Carr *
Fenland (disambiguation) Fenland may mean: * Fenland, or the Fens, an area of low-lying land in eastern England ** Fenland District, a local authority district in Isle of Ely, Cambridgeshire, England, forming part of the Fens ** Fenland Airfield, an airfield near Spalding, ...
* Fen-meadow *
Reed bed A reedbed or reed bed is a natural habitat found in floodplains, waterlogged depressions and estuaries. Reedbeds are part of a succession from young reeds colonising open water or wet ground through a gradation of increasingly dry ground. As ...


Specific fens

* Wicken Fen, Cambridgeshire *
The Fens The Fens, also known as the , in eastern England are a naturally marshy region supporting a rich ecology and numerous species. Most of the fens were drained centuries ago, resulting in a flat, dry, low-lying agricultural region supported by a ...
(eastern England) *
Mesopotamian Marshes The Mesopotamian Marshes, also known as the Iraqi Marshes, are a wetland area located in Southern Iraq and in southwestern Iran. The marshes are primarily located on the floodplains of the Euphrates and Tigris rivers bound by the cities of Basra, ...
(Iraq) * Back Bay Fens (Boston, Massachusetts) *
Cedar Bog Cedar Bog State Nature Preserve is a fen left behind by the retreating glaciers of the Wisconsin glaciation about 12,000-18,000 years ago. A protected area of about of fen remains from the original area of approximately 7,000 acres (28 km2) ...
(Champaign County, Ohio) *
Cowles Bog Cowles Bog is a 4,000-year-old wetland complex in Indiana Dunes National Park, near Chesterton, Indiana. It is named for Henry Chandler Cowles who did his pioneering work in ecology and ecological succession here. His work brought international a ...
, Indiana Dunes National Lakeshore (Indiana, USA) *
Geneva Creek (Colorado) Geneva Creek is a short tributary of the North Fork South Platte River, approximately long,U.S. Geological Survey. National Hydrography Dataset high-resolution flowline dataThe National Map , accessed March 25, 2011 in central Colorado in the Uni ...
(an iron fen in the USA)


References


Citations


General bibliography

* * * * * * * *


External links

* {{Authority control Landforms Limnology Soil Wetlands