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• Wetland Ecology
• Lectures 14-15-16
• Wetland Biogeochemistry
• What is biogeochemical cycling?
– Transport & Transformation of chemicals in an ecosystem, involving numerous interrelated physical, chemical, & biological processes
• Examples of movement include
– Water-sediment exchange
– Plant Uptake
– Organic exports
• Two major categories of wetland biogeochemistry include:
– Intrasystem cycling through various transformation processes
– Exchange of chemicals between a wetland & its surroundings
• Open vs Closed Ecosystems
OPEN
• BHF + Tidal Marshes have significant exchange of materials with surroundings
– River flooding + Tidal Exchange
CLOSED
• Bogs + Cypress Domes
– Little material exchange except for gaseous matter!
• Biogeochemically open vs closed
• BGC Open – Abundant exchange of materials with surroundings
– Tidally driven or riparian
• BGC Closed – Little movement of materials across the ecosystem boundary
– Cypress swamps or very stagnant areas
• So, are wetlands open or closed systems?
– They can be both!
• Wetland Soil
• Wetland soil is:
– Medium in which many of the wetland chemical transformations occur
– The primary storage of available chemicals for wetland plants
• Two major types of wetland soils
– Organic & Organic Mineral Soils
– Mineral
• Organic Soil + Organic Mineral Soils
Defined under two saturation conditions
• 1. Saturated with water for long periods
– Have > 18% organic carbon
– Have > 12% organic carbon if no clay present
– Have a proportional content of organic carbon (12 – 18%) if clay content (0 – 60%)
• 2. Soils are never saturated with water for more than a few days and have > 20% organic carbon
– %Corg = %OM/2
– %Corg = percentage of organic carbon
– %OM = percentage of organic matter
• Bulk density & porosity – have lower bulk densities (dry weight of soil/volume). Low due to high porosity (peat soils ~ 80% air space)
• Hydraulic conductivity – Depends on degree of decomposition. Organic soils hold more water (do not necessarily allow more water to pass)
• Nutrient availability – Organic soils have more minerals tied up in organic form.
• Cation Exchange Capacity – Organic soils have greater CEC (sum of exchangeable ions)
• Organic soils (What order?) are classified into four groups; 3 are hydric
– Saprists (muck) – > 2/3 of the material is decomposed, < 1/3 of plant fibers are identifiable
– Fibrists (peat) - < 1/3 of material is decomposed and > 2/3 of plant fibers are identifiable
– Hemists (mucky peat or peaty muck) – Conditions fall between saprist & fibrist soil
– Folists – organic soils caused by excessive moisture (precip > evapotranspiration) that accumulate in tropical & boreal mountains; not classified as hydric because saturated conditions are the exception rather than the rule
• Mineral Wetland Soil
• When flooded for extended periods mineral soils develop certain characteristics that allow for their identification
– Redoximorphic features (mediated by microbes)
– The rate these features are formed depend on three conditions (all must be present)
• Sustained anaerobic conditions
• Sufficient soil temp (5°C “biological zero”)
• Organic Matter (substrate for microbial activity)
• Hydric mineral soils are characterized by:
– Gleying
– Oxidized Rhizosphere
– Mottles (aka Redox Concentrations)
– Current nomenclature
• 1. Redox concentrations – Accumulation of Fe & Mn in 3 different structures
– Nodules & Concretions (firm extremely firm irregularly shaped bodies with diffuse boundaries)
– Masses – formerly called ‘reddish mottles’
– Pore linings – Formerly included ‘oxidized rhizospheres’
• 2. Redox depletions – Low-chroma (<2) bodies with high values (>4) including:
– Iron depletions: “Gray mottles” or “Gley mottles”
– Clay depletions: Contain less Fe, Mn, and clay than adjacent soils
• 3. Reduced matrices: Low-chroma soils
• REDOX in Wetlands
• When mineral or organic soils are flooded anaerobic conditions result. Water fills pore spaces and rate of oxygen diffusion through soil is drastically reduced
• Rate of oxygen depletion depends on:
– Ambient Temperature
– Availability of organic substrates for microbial respiration
– Chemical oxygen demand from reductants such as ferrous Fe
• Resultant O2 deficiency prevents plants from normal aerobic root respiration and affect nutrient availability and adds toxic materials in the soil
• Usually thin layer formed and is related to:
– Rate of O2 transport across the atmosphere-surface water interface
– Small population of O2 consuming organisms
– Photosynthetic O2 production by algae within water column
– Surface mixing by convection currents & wind action
• Eh ranges
• If DO is present, the redox potential range +400 to + 700mV
• If O2 disappears, Eh range from +400 down to -400mV
• As organic substrates in a water logged soil are oxidized (donating) the redox potential drops = a sequence of reductions (gains) takes place
• 1st reaction to occur after becoming anaerobic is the reduction of NO3- (nitrate) first to NO2-
(nitrite) and ultimately to N2O or N2
– Nitrate becomes an acceptor ~ 250mV
– At 225mV Mn is transformed from manganic to manganous
– At -75 to -150mV Fe is transformed from ferric to ferrous; while sulfates are reduced to sulfides
• pH
• Soil and overlying waters of wetlands occur over a wide range of pH’s
– Organic soils – often more acidic
– Mineral soils – often neutral or alkaline
• Alkaline soils previously drained decrease in pH because of buildup of CO2 then carbonic acid
• Acid soils previously drained increase in pH because of reduction of ferric iron hydroxide
• Nitrogen Transformations
• Nitrogen is often the “most limiting nutrient in flooded soils”
– Limitations reported in salt marshes, freshwater inland marshes, & freshwater tidal marshes
• Involve complex microbial processes
• NH4 is the primary form of mineralized N in wetlands
• Mineralization
• Often referred to as ammonification
NH2CONH2 + H2O 2NH3 + CO2
NH3 + H2O NH4 + OH-
• Nitrification
• Once ammonia has formed, it can take several possible pathways
– Aerobic environment – ammonium is oxidized (nitrification) in two steps by Nitrosomonas
sp.
2NH4+ + 3O2 2NO2
- + 2H2O + 4H
+ + energy
– By Nitrobacter sp.
2NO2- + O2 2NO3
- + energy
• Denitrification
• Denitrification is carried out by facultative bacteria under anaerobic conditions – nitrate is the terminal electron acceptor
C6H12O6 + 4NO3 6CO2 + 6H2O + 2N2
• Most significant path of nitrogen loss from wetlands
• Usually lost as N2 & N2O
• N fixation
• Conversion of N2 gas to organic nitrogen
– Favored by low oxygen concentrations
– Rhizobium species
– Cyanobacteria (blue-green algae) are also common in Louisiana, northern bogs, & in rice cultures
– Fe & Mn Transformations
• Found in reduced forms in wetlands
• Readily available Toxic levels
• More soluble
– Ferrous Fe reduced form of Fe
– Ferric Fe oxidized form of Fe
• Oxidized form creates barrier; may prevent plant from uptaking other nutrients
• Reduced form reacts with P making it unavailable
• Sulfur
• Rarely limiting to plant or animal growth in wetlands
– Hydrogen sulfide (H2S) is toxic (rotten-egg smell)
– Ferrous sulfide is responsible for black color of wetland soils (highly reduced sediment)
• Negative effects of sulfides on higher plants are attributable to a number of causes
– Direct toxicity of free sulfide as it comes into contact with plant roots
– Reduced availability of sulfur for plant growth because of its precipitation with trace metals
– Immobilization of zinc & copper by sulfide precipitation
• Carbon Cycle
• Photosynthesis & aerobic respiration dominate the aerobic horizons
• Fermentation
• Methanogenesis
• Occurs when certain bacteria (methanogens) use CO2 as an electron acceptor for the production of gaseous methane (CH4)
• Requires extremely reduced conditions
– Redox potential range from -250 to -350mV
• Phosphorus
• One of the most important chemicals in wetland systems
• Most limiting in northern bogs, freshwater marshes, & southern deepwater swamps
• Retention is one of the most important features for natural & constructed wetlands
• Principle inorganic form = orthophosphate
– PO43-
(pH > 13)
– H2PO4- (pH 2-7)
– HPO4—
(pH 8-12)
• Predominant form dependent on pH
• P is not directly altered by Eh changes
– Affected by association with other elements especially Fe
• P is rendered relatively unavailable to plants & microconsumers by:
– 1. Precipitation of insoluble phosphates with ferric Fe, Ca, & Al under aerobic conditions
– 2. Adsorption (particle surface) of phosphate onto clay particles, organic peat and ferric & aluminum hydroxides & oxides
– 3. Binding of phosphorous in OM as a result of its incorporation into the living biomass of bacteria, algae, & vascular macrophytes
• Chemical Transport into Wetlands
• Geologic inputs – from weathering of rock
– Type of soil direct reflection of parent material
• Biologic inputs – Photosynthetic uptake of carbon, nitrogen fixation, & biotic transport of materials by animals (birds)
• Hydrologic inputs – Major inputs into wetlands
• Precipitation
– Burning of fossil fuels
– Increased [ ]’s of sulfates & nitrates in atmosphere
• Streams, Rivers, & Groundwater
– As precip reaches the ground it will:
– Infiltrate into the ground
– Return to atmosphere via evapotranspiration
– Flow on surface as runoff
• Groundwater influence:
– Chemical characteristics of streams & rivers depend on the degree to which the water has previously come into contact with underground formations & types of minerals present in those formations
• Climate:
– Balance of precipitation & evapotranspiration
– Type of vegetation present
• Geographic effects: amount of dissolved & suspended materials that enter streams, rivers, & wetlands depend on:
– Size of watershed
– Steepness or slop of landscape
– Soil texture
• Streamflow/Ecosystem effects: The water quality of surface water runoff, streams, & rivers varies seasonally
• Human effects: water that has been modified by humans through sewage effluent, urbanization, & runoff from farms alters the chemical composition of streamflow & groundwater that enters wetlands
• Drainage from agriculture fields:
– Higher [ ]’s of sediments, nutrients, herbicides, & pesticides might be expected
• Drainage from urban & suburban areas:
– May have high [ ]’s of trace organics, oxygen demanding substances, & some toxic materials
• Estuaries
• Quality differs from that of rivers
• Seawater chemical composition is fairly constant worldwide
– 33 0/00 to 37 0/00
– Mass balances
• A quantitative account of the inputs, outputs, & internal cycling of materials in an ecosystem
• Mass balances help determine:
– Ecosystem functions
– Determine the importance of wetlands as sources, sinks, and transformers of chemicals
• Inputs primarily through:
– Hydrologic
• Precipitation
• Surface & groundwater inflow
• Tidal Exchange
– Biotic
• Atmospheric carbon fixation (photosynthesis)
• Atmospheric nitrogen (nitrogen fixation)
• Exports:
– Surface water & groundwater
– Long-term burial of chemical in the sediments
• Intrasystem cycling involves exchanges among various pools, or standing stocks of chemicals in within a wetland. Involves pathways such as:
– Litter production
– Remineralization
– Chemical transformations
– Translocation of nutrients through plants
• Generalizations
• 1. Wetlands serve as sources, sinks, or transformers of chemicals, depending on the wetland type, hydrologic condition, & length of time the wetland has been subjected to chemical loadings
• 2. Seasonal patterns of nutrient uptake & release
– Temperate regions retention is higher in growing season
• Increased microbial activity
• Higher macrophyte activity
• 3. Wetlands are frequently coupled to adjacent ecosystems through chemical exchanges that affect both systems
– Downstream ecosystems benefit from retention or from exportation
• 4. Wetlands are either highly productive (eutrophic) or low productivity (oligotrophic)
• 5. Nutrient cycling in wetlands differs from aquatic & terrestrial systems
– More nutrients in sediment & peat
– Aquatic systems have autotrophic activity more dependent on nutrients in water column than in sediments.
– Wetland plants obtain nutrients from sediment
• 6. Anthropogenic changes have led to changes in nutrient cycling in many wetlands
• The capacity of wetlands to assimilate anthropogenic wastes from the atmosphere or hydrosphere is not unlimited!