Difference between revisions of "Estuarine ecosystems"

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==Introduction==
 
==Introduction==
  
[[Estuary|Estuaries]], or transitional waters, represent the transition between freshwater and marine environments and are influenced by both aquatic realms. Salinity levels are indicative of the position within the mixing zones of an estuary. The upper limit of an estuary is referred to as its head, while the lower limit is called the mouth of the estuary. Between the freshwater head and the saline mouth of the estuary lie a number of zones marked by intermediate salinity values, each with distinct characteristics pertaining to light penetration and type of substratum, thus hosting different communities of organisms.
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[[File:EstuaryHighLowWater.jpg|thumb|right|500px|Fig. 1. Schematic planview of a typical estuarine morphology at high water (HW, top) and low water (LW, bottom). ]] 
  
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[[Estuary|Estuaries]], or transitional waters, represent the transition between freshwater and marine environments and are influenced by both aquatic realms. Salinity levels are indicative of the position within the mixing zones of an estuary. The upper limit of an estuary is referred to as its head, while the lower limit is called the mouth of the estuary. Between the freshwater head and the saline mouth of the estuary lie several zones corresponding to intermediate salinity ranges (Fig. 1). These zones also differ in other respects, such as light penetration and the type of substrate, and therefore host different communities of organisms. However, it should be realized that these zones move up and down the estuary with the tide over distances typically on the order of 10 - 20 km. Swimming and floating organisms move with the tide, but benthic organisms do not. They have to cope with tidally modulated salt contents that can vary between oligohaline and mesohaline or between mesohaline and polyhaline. On the time scale from month to season, the variation in salinity is even greater because of variations in river discharge.
  
[[Image:Estuarine division.jpg|thumb|centre|500px|Division of transitional waters (Carricker, 1967<ref> Carriker, M. R. (1967). Ecology of estuarine benthic invertebrates: a perspective, p. 442-487. In G. H. Lauff (ed.), Estuaries, American Association for the Advancement of Science. Washington. </ref>; McLusky, 1989<ref> McLusky, D. S. (1989). The Estuarine Ecosystem, 2nd Ed. Chapman & Hall, London. </ref>)]]
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{| style="border-collapse:collapse;background:ivory; font-size: 12px " cellpadding=5px align=center width=90%
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|+ Division of transitional waters for typical estuaries. Adapted from  Carriker, 1967<ref>Carriker, M. R. 1967. Ecology of estuarine benthic invertebrates: a perspective, p. 442-487. In G. H. Lauff (ed.), Estuaries, American Association for the Advancement of Science. Washington.</ref>; McLusky, 1989<ref>McLusky, D. S. 1989. The Estuarine Ecosystem, 2nd Ed. Chapman & Hall, London.</ref>.
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|- style="font-weight:bold; font-size: 11px; text-align:center; background:lightblue"
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! width="20%" style=" border:1px solid blue; font-size: 11px;"| Estuary division
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! width="10%" style=" border:1px solid blue; font-size: 11px;"| Length scale [km]
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! width="10%" style=" border:1px solid blue; font-size: 11px;"| Subtidal (channel) substratum
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! width="10%" style=" border:1px solid blue; font-size: 11px;"| Intertidal (mudflat, saltmarsh) substratum
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! width="10%" style=" border:1px solid blue; font-size: 11px;"| Salinity range
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! width="10%" style=" border:1px solid blue; font-size: 11px;"| Zone
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! width="20%" style=" border:1px solid blue; font-size: 11px;"| Organism type
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|-
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| style="border:2px solid lightblue; font-size: 11px; text-align:center"| river
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| style="border:2px solid lightblue; font-size: 11px; text-align:center"| -
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| gravel / sand
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| -
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| < 0.05
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| limnetic
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| freshwater
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|-
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| tidal river (seaward of tidal limit)
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 10 - 100
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| sand
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| silt / mud
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 0.05 - 0.5
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| limnetic
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| freshwater
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|-
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| estuary head (seaward of salt intrusion limit)
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 2 - 5
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| sand / silt
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| silt / mud
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 0.5 - 5
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| oligohaline (brackish)
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| oligohaline, freshwater migrants
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|-
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| upper reach
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 5 - 25
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| sand /silt / mud
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| silt / mud
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 5 - 18
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| mesohaline
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| true estuarine, limit of non-transient migrants
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|-
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| middle reach
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 5 - 25
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| sand / silt / mud
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| silt / mud
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 18 - 25
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| polyhaline
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| estuarine, euryhaline
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|-
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| lower reach
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 5 - 10
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| sand
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| silt / mud
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| 25 – 30
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| polyhaline
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| style="border:2px solid lightblue; font-size: 11px;text-align:center"| estuarine, euryhaline, marine migrants
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|-
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| style="border:2px solid lightblue; font-size: 12px;text-align:center"| mouth / sea
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| style="border:2px solid lightblue; font-size: 12px;text-align:center"| 5 - 10
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| style="border:2px solid lightblue; font-size: 12px;text-align:center"| sand / gravel / rock
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| style="border:2px solid lightblue; font-size: 12px;text-align:center"| -
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| style="border:2px solid lightblue; font-size: 12px;text-align:center"| 30-35
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| style="border:2px solid lightblue; font-size: 12px;text-align:center"| euhaline
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| style="border:2px solid lightblue; font-size: 12px;text-align:center"| stenohaline, all marine
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|}
  
  
 
== The challenges of estuarine ecosystems ==
 
== The challenges of estuarine ecosystems ==
  
Estuaries are peculiar yet challenging ecosystems. The main challenge and at the same time the most important feature governing species diversity of transitional waters is the '''variable salinity regime'''. Salt dissolved in water dehydrates living organisms by exerting what is called osmotic pressure on the cell walls. Organisms living in the sea are equipped with buffering mechanisms allowing them to retain their body fluids in the presence of salt. In most cases this mechanism is adjusted to a particular salinity and yields a distinction between freshwater and marine species. Most aquatic organisms fall into one of these categories. In turn very few species can withstand variable salinity, which has implications for the biodiversity of estuaries.  
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[[File:MudflatOrganisms.jpg|thumb|right|300px|Fig. 2. Typical mudflat organisms.]]
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Organisms that live in estuaries must be adapted to the specific conditions that occur here. The most constraining condition is the varying salinity. Most organisms are so-called '''stenohalines''': they survive only in an environment with a certain more or less constant salinity, for example in the sea or in rivers. These species cannot survive in an estuarine environment. Species that thrive in an estuarine environment must be able to cope with varying [[Osmosis|osmotic pressures]] related to varying salinity; these species are called '''euryhalines'''. When the internal osmotic pressure of a cell is lower than the external pressure, the cell will lose fluid and shrink; in the opposite case, the cell will swell. There are two strategies to counter this. One strategy is used by the so-called [[Osmosis#Osmoconformers|osmoconformers]]: they regulate the internal osmotic pressure in such a way that it becomes equal to the external osmotic pressure. Depending on the ambient salinity, they achieve this by producing the required amount of organic osmolytes (molecules such as sugars, amino acids, methylamines) to balance the internal and external osmotic pressure<ref>Yancey, P.H. 2005. Organic osmolytes as compatible, metabolic and counteracting cytoprotectants in high osmolarity and other stresses. J. Exp. Biol. 208: 2819–2830</ref>. The other strategy is used by the so-called [[Osmosis#Osmoregulators|osmoregulators]]: they regulate the osmotic pressure by secreting an excess of salt or an excess of water, depending on the salinity of the environment. Because euryhaline species are relatively rare, the estuarine ecosystem is characterized by a relatively limited biodiversity (Fig. 2) but a high abundance of specific estuarine species.
  
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Most marine invertebrates are osmoconformers and generally stenohaline. However, some species are euryhaline, for example barnacles, some polychaetes worms and some copepods. Bivalve mollusks are euryhaline osmoconformers within a limited salinity range, but cope with strong salinity variations by closing their valves. Marine vertebrates are generally osmoregulators. Several crustacean species (e.g. crabs) and fish species (e.g. salmon, flounder) are euryhaline. Euryhaline osmoregulating vegetation species include ''Spartina'' and ''Salicornia''. Mangroves, however, are adapted to specific salinity ranges and thus cannot be classified as euryhaline.
  
  
[[Image:Shannon_Estuary.jpg|right|thumb|350px|caption|Turbid waters of the Shannon Estuary]]
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Lowland reaches of rivers are characterised by high levels of suspended solids, such as silt, clay and organic [[detritus]], inducing high [[Turbidity|turbidity]]. In the fresh water – sea water transition zone these particles become effectively ‘trapped’ due to [[Flocculation cohesive sediments|flocculation]] and converging suspended sediment fluxes <ref>Kranck, K. (1981) Particulate matter grain-size characteristics and flocculation in a partially mixed estuary. Sedimentology 28: 107 – 114.</ref><ref> Robinson M. C., Morris, K. P. & Dyer, K. R. (1999). Deriving fluxes of suspended particulate matter in the Humber Estuary, UK, using airborne remote sensing. Marine Pollution Bulletin, 37: 155 -163</ref>. These convergences result in a so-called [[estuarine turbidity maximum]] (ETM). For further explanation, see the articles [[Estuarine turbidity maximum]] and [[Dynamics of mud transport]].
  
Lowland reaches of rivers are characterised by high levels of suspended solids, such as silt, clay and organic [[detritus]], inducing high [[Turbidity|turbidity]]. In the fresh water – sea water transition zone these particles become effectively ‘trapped’ due to [[flocculation]] and converging suspended sediment fluxes <ref>Kranck, K. (1981) Particulate matter grain-size characteristics and flocculation in a partially mixed estuary. Sedimentology 28: 107 – 114.</ref><ref> Robinson M. C., Morris, K. P. & Dyer, K. R. (1999). Deriving fluxes of suspended particulate matter in the Humber Estuary, UK, using airborne remote sensing. Marine Pollution Bulletin, 37: 155 -163.</ref> (see also [[Dynamics of mud transport]]). These convergences result in pronounced estuarine turbidity maxima (ETM).
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Turbidity limits the depth of the photic zone (light penetration zone), thus limiting the photosynthesis and primary production. Much of the primary production in estuaries occurs on the seaward side of the ETM (Swart, 2007<ref> Swart, de H. E., Schuttelaars, H. M. & Talke, S. A. (2007). A simple model for phytoplankton growth in turbid estuaries. Geophysical Research Abstracts 9, 04190. </ref>). Macrophyte vegetation and benthic algae are often limited to the periodically exposed (intertidal) part of the estuary, while the growth of phytoplankton is restricted to a thin uppermost layer of the water column (Cloern, 1997<ref> Cloern, J.E. (1997). Turbidity as a control on phytoplankton biomass and productivity in estuaries. Continental Shelf Research 7: 1367-1381.</ref>.) As a result, estuaries are '''heterotrophic systems''', where more energy is consumed than produced. Most estuarine species are detritivores, meeting their energy intake requirements from organic matter contained either in the sediment (deposit feeders), in suspension (filter feeders) or both. Benthic invertebrates play a major role in energy cycling in estuaries (Wilson & Parkes, 1998<ref> Wilson, J.G. & Parkes, A. (1998). Network analysis of the energy flow through the Dublin Bay ecosystem. Biology And Environment: Proceedings Of The Royal Irish Academy 98B (3): 179–190. </ref>).
  
Turbidity limits the depth of the photic zone (light penetration zone), thus limiting the photosynthesis and primary production. Much of the primary production in estuaries occurs on the seaward side of the ETM (Swart, 2007<ref> Swart, de H. E., Schuttelaars, H. M. & Talke, S. A. (2007). A simple model for phytoplankton growth in turbid estuaries. Geophysical Research Abstracts 9, 04190. </ref>). Macrophyte vegetation and benthic algae are often limited to the periodically exposed (intertidal) part of the estuary, while the growth of phytoplankton is restricted to a thin uppermost layer of the water column (Cloern, 1997<ref> Cloern, J.E. (1997). Turbidity as a control on phytoplankton biomass and productivity in estuaries. Continental Shelf Research 7: 1367-1381.</ref>.) As a result, estuaries are '''heterotrophic systems''', where more energy is consumed than produced. Most estuarine species are detritivores, meeting their energy intake requirements from organic matter contained either in the sediment (deposit feeders), in suspension (filter feeders) or both. Benthic invertebrates play a major role in energy transfer and circulation in estuaries (Wilson & Parkes, 1998<ref> Wilson, J.G. & Parkes, A. (1998). Network analysis of the energy flow through the Dublin Bay ecosystem. Biology And Environment: Proceedings Of The Royal Irish Academy 98B (3): 179–190. </ref>).
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[[File:DebenEstuary.JPG|right|thumb|350px|caption|Fig. 3. Mudflat in the Deben estuary (UK).]]
  
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Particulate matter eventually settles in the estuarine mid reaches where the substratum can contain a high mud fraction, especially in the intertidal areas adjacent to the main tidal channels (Fig. 3). These mudflats are less suitable for organisms with an affinity for hard substratum but provide an important habitat for a range of burrowing invertebrate species, of which bivalve mollusks and polychaete worms are usually dominant in terms of numbers and biomass. These in turn provide an attractive dish for higher trophic consumers.
  
Particulate matter eventually settles out and the substratum of estuarine mid reaches tends to be composed of fine fractions, typically creating '''extensive sand and mudflats'''. These conditions hinder organisms with an affinity for hard substratum but constitute important habitats for a range of burrowing invertebrate species, of which bivalve molluscs and polychaete worms are usually dominant in terms of numbers and biomass. These in turn provide a rich feeding opportunity for a range of higher consumers.  
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Sediment deposition gradually increases the height of the intertidal areas until submergence only occurs at the highest tides. Accretion is stimulated by the development of vegetation on the landward zones which are most sheltered from currents and waves. These high vegetated zones of the intertidal area, the so-called saltmarshes, form an important estuarine habitat. This habitat is described in the article [[Salt marshes]].
  
  
Industrialised and urbanised river catchments and estuaries receive anthropogenic input from various sources (both point sources and diffuse sources) and often carry a range of contaminants. Sheltered, low-energy areas such as intertidal mudflats in enclosed bays or estuaries are most susceptible to these pollutants due to slow dispersion. The finer substrata in these areas act as a '''sink for contaminants''' making estuarine benthic fauna susceptible to pollution (Kausch & Michaelis, 1996<ref> Kausch, H. & Michaelis, W. (eds.) (1996).  Suspended particulate matter in rivers and estuaries - Proceedings of an International Symposium held at Reinbek near Hamburg, Germany. Advances in Limnology, 47. </ref>).
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Industrialised and urbanised river catchments and estuaries receive anthropogenic input from various sources (both point sources and diffuse sources) and often carry a range of contaminants. These pollutants bind to fine sediments and therefore can accumulate in sheltered, low-energy areas such as intertidal mudflats in enclosed bays or estuaries. The estuarine benthic fauna of these zones can thus be subject to high contaminant levels (Kausch and Michaelis, 1996<ref>Kausch, H. and Michaelis, W., eds. 1996.  Suspended particulate matter in rivers and estuaries - Proceedings of an International Symposium held at Reinbek near Hamburg, Germany. Advances in Limnology, 47. </ref>). For further details, see: [[Coastal and marine sediments]].
  
See also: [[Coastal and marine sediments]]
 
  
 
== Biodiversity of estuarine ecosystems ==
 
== Biodiversity of estuarine ecosystems ==
  
[[Image:Remane diagram.jpg|thumb|left|400px|Species richness in estuaries follows a long established pattern along the salinity gradient described by Remane (1934<ref> Remane, A. (1934). Die Brackwasserfauna. Verzeichnis der Veröffentlichungen Goldsteins, 36: 34–74.
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[[Image:Remane diagram.jpg|thumb|left|250px|Fig. 4. Species richness in estuaries follows a long established pattern along the salinity gradient described by Remane (1934<ref> Remane, A. (1934). Die Brackwasserfauna. Verzeichnis der Veröffentlichungen Goldsteins, 36: 34–74.
 
.</ref>)]]
 
.</ref>)]]
  
Truly '''estuarine species''' are those that complete their whole life cycle within the transitional waters. Species permanently dwelling there are mostly hardy, stress-tolerant species able to handle salinity shifts and high suspended solid levels. Not many species can perform well under such conditions. Estuarine ecosystems are thus typically characterised by relatively low species diversity comparing to freshwater or full salinity conditions. Freshwater species are becoming less abundant with increasing salinity and are gradually replaced by marine organisms moving down the estuary with some truly estuarine species found at intermediate salinities. This pattern is reflected by the overall species richness, where the least diverse fauna is found at salinity levels of between 5 and 18 PSU.
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Truly estuarine species are those that complete their whole life cycle within the transitional waters. Species permanently dwelling there are mostly hardy, stress-tolerant species able to handle salinity fluctuations and high suspended solid levels. Organisms living on the higher parts of the mudflats and on the salt marshes are subject to additional stresses during temporary emersion at low tide, such as exposure to air, dehydration and temperature variations. Not many species can perform well under such conditions. Estuarine ecosystems are thus characterised by relatively low species diversity compared to freshwater or full salinity conditions. Along the estuary, from head to mouth, freshwater species become rare as salinity increases and are gradually replaced by marine organisms in the lower estuarine reaches, with some truly estuarine species found at intermediate salinities. This pattern is reflected by the overall species richness, where the least diverse fauna is found at salinity levels between 5 and 18 ppt. This is reflected in the so-called Remane curve, see Fig. 4.  
  
  
Apart from the permanent dwellers, estuaries are host to a number of '''visitors'''. Some of them have to travel through estuaries on their migratory route, being either anadromous (spawn in freshwater and feed and grow at sea) or catadromous (spawn at sea and feed and grow in freshwater). The absence of many of the marine predators and rich particulate food supply offer attractive spawning and nursery grounds for many species that normally live under full salinity conditions. Even though estuarine ecosystems are usually species-poor, they maintain a high productive throughput of invertebrate fauna. This productivity provides rich feeding opportunities for a range of higher consumers. Some marine predators are well equipped to cope with reduced salinity and frequently enter estuaries in search of food. Ebb tide makes estuarine beds available to terrestrial predators, of which birds take the greatest share. Estuarine sand and mudflats that are periodically exposed to air support high densities of a diverse bird fauna. This value of estuaries has been long recognized and many estuarine sand and mudflats have been designated as Special Protection Areas (SPA) under the EU Birds Directive (79/409/EEC).
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Apart from the permanent dwellers, estuaries are host to a number of visitors. Some of them have to travel through estuaries on their migratory route, being either anadromous (spawn in freshwater and feed and grow at sea) or catadromous (spawn at sea and feed and grow in freshwater). The absence of many of the marine predators and rich particulate food supply offer attractive spawning and nursery grounds for many species that normally live under full salinity conditions. Even though estuarine ecosystems are usually species-poor, they maintain a high productive throughput of invertebrate fauna. This productivity provides rich feeding opportunities for a range of higher consumers. Some marine predators are well equipped to cope with reduced salinity and frequently enter estuaries in search of food. Ebb tide makes estuarine beds available to terrestrial predators, of which birds take the greatest share. Estuarine sand and mudflats that are periodically exposed to air support high densities of a diverse bird fauna. This value of estuaries has been long recognized and many estuarine sand and mudflats have been designated as [[Birds Directive, Habitats Directive, NATURA 2000|Special Protection Areas (SPA)]] under the EU Birds Directive (79/409/EEC).
  
  
 
==Threats==
 
==Threats==
  
Estuarine ecosystems are usually dominated by stress-tolerant organism, able to withstand a relatively wide range of environmental factors. However, they to face some threats from anthropogenic activities.  
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Estuarine ecosystems are usually dominated by stress-tolerant organism, able to withstand a wide range of environmental conditions. However, they also have to face serious threats from anthropogenic activities.
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[[File:ChannelizationLoireEstuary.jpg|right|thumb|350px|Caption|Fig. 5. Historical maps of the Loire estuary channelization. Photo: Groyne field along the northern channel bend (at the right in the maps). ]]
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* Estuaries are preferred locations for human settlement, especially due to the natural connections they offer between inland and overseas destinations. Residential, recreational and industrial developments (such as harbours or ports and marinas) are usually located right at the waterfront with supporting structures (roads, railways) that cut off the connections with the upper shore habitats. Estuaries  are often challenged by land development; land reclamation is particularly detrimental in this respect as it results in a permanent loss of habitat.
  
* Estuaries are preferred locations for human settlement due to their high productivity and availability of natural connections between maritime and inland waterways. Residential, recreational and industrial developments (such as marinas, harbours or ports) are usually located right at the waterfront with supporting structures such as embankments impacting on the upper shore communities. Estuaries  are often challenged by '''land development'''; land reclamation is particularly detrimental in this respect as it results in a permanent loss of habitat.
 
  
[[Image:Artificial embankments.jpg|right|thumb|500px|Caption|Highly polluted and channeled section of the Liffey Estuary.]]
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* Making estuaries with shifting shoals and channels navigable has long been beyond human capabilities. This changed at the end of the 19th century, when large-scale dredging technology became available. Without exception, all estuaries on which harbors were located, have been channeled in the 20th century (Fig. 5). A deep navigation channel was dredged, often deviating from the natural tidal channel. The channel banks were set in stone and groynes were constructed to concentrate the flow and to suppress the natural tendency of tidal channels to shift and meander. Channel deepening allowed the tide to propagate further up the river (with especially a strong lowering in the low water level) and both salt intrusion and turbidity increased, sometimes to a very large extent. Fixing the channel banks has led to strong sedimentation of the remaining intertidal areas, causing the mud flat and salt marsh levels to rise. The raised salt marshes have often been embanked and turned into agricultural land or have become terrestrial habitats. The dynamics of migrating tidal channels, the natural agent of habitat rejuvenation and new marsh development, has therefore been thwarted and replaced by a development towards aging landscapes with significant loss of biodiversity.
  
  
* Rivers discharging into their estuaries carry various constituents depending on  the landuse of the drainage area (catchment). This means that various contaminants introduced at any point in the catchment ultimately end up in the estuary. Although estuarine organisms are typically hardy, these '''pollutants and excess nutrients''' impede their overall performance (including growth and reproduction). Pollution from densely populated or heavily industralised catchments has detrimental effects on life in estuaries. See: [[Eutrophication in coastal environments]], [[Coastal pollution and impacts]].
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* Rivers discharging into their estuaries carry various constituents depending on  the landuse of the drainage area (catchment). This means that various contaminants introduced at any point in the catchment ultimately end up in the estuary. Although estuarine organisms are typically hardy, these pollutants and excess nutrients impede their overall performance (including growth and reproduction). Pollution from densely populated or heavily industralised catchments has detrimental effects on life in estuaries. See: [[Eutrophication in coastal environments]], [[Coastal pollution and impacts]].
  
  
* Climate change impacts of concern for estuaries are the overall temperature rise and elevation of the sea level. The first one is likely to induce latitudinal migration with more warm-water species being increasingly established and possibly outcompeting the native species in a long run. Sea level rise would result in a shift of the salinity zonation landwards (see also [[Effects of global climate change on European marine biodiversity]]). However, coastal areas, including estuaries, are usually heavily populated and developed. In such developed areas the vertical shift of salinity zonation can be hindered by flood defence structures and river and shore embankments resulting in estuarine squeeze.
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* Climate change impacts of concern for estuaries are the overall temperature rise and elevation of the sea level. The former is likely to induce latitudinal migration with more warm-water species being increasingly established and possibly outcompeting the native species in a long run. Sea level rise would result in a shift of the salinity zonation landwards (see also [[Effects of global climate change on European marine biodiversity]]). However, coastal areas, including estuaries, are usually heavily populated and developed. In such developed areas the vertical shift of salinity zonation will be hindered by flood defence structures and river and shore embankments resulting in estuarine [[Coastal squeeze|squeeze]]. Because many estuaries are situated in lowland, a substantial rise of the high water levels will have great consequences for the safety of the people living in these low-lying areas and for their properties and infrastructure. See the article [[Coastal cities and sea level rise]].  
  
  
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==Related articles==
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:[[Morphology of estuaries]]
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:[[Estuarine turbidity maximum]]
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:[[Salt marshes]]
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:[[Mangroves]]
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:[[Blue carbon sequestration]]
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:[[Tidal asymmetry and tidal basin morphodynamics]]
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:[[Coastal and marine sediments]] 
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:[[Estuarine morphological modelling]]
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:[[Osmosis]]
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:Other article in the category [[:Category:Estuaries and tidal rivers|Estuaries and tidal rivers]]
  
== Further reading ==
 
* Estuarine and Coastal Sciences Association https://ecsa.international/
 
* Joint Nature Conservation Comitee (JNCC) - Estuaries http://www.jncc.gov.uk/page-2160
 
* US Environmental Protection Agency - Estuarine Science https://www.epa.gov/nep
 
  
  

Revision as of 21:55, 17 August 2021



Introduction

Fig. 1. Schematic planview of a typical estuarine morphology at high water (HW, top) and low water (LW, bottom).

Estuaries, or transitional waters, represent the transition between freshwater and marine environments and are influenced by both aquatic realms. Salinity levels are indicative of the position within the mixing zones of an estuary. The upper limit of an estuary is referred to as its head, while the lower limit is called the mouth of the estuary. Between the freshwater head and the saline mouth of the estuary lie several zones corresponding to intermediate salinity ranges (Fig. 1). These zones also differ in other respects, such as light penetration and the type of substrate, and therefore host different communities of organisms. However, it should be realized that these zones move up and down the estuary with the tide over distances typically on the order of 10 - 20 km. Swimming and floating organisms move with the tide, but benthic organisms do not. They have to cope with tidally modulated salt contents that can vary between oligohaline and mesohaline or between mesohaline and polyhaline. On the time scale from month to season, the variation in salinity is even greater because of variations in river discharge.


Division of transitional waters for typical estuaries. Adapted from Carriker, 1967[1]; McLusky, 1989[2].
Estuary division Length scale [km] Subtidal (channel) substratum Intertidal (mudflat, saltmarsh) substratum Salinity range Zone Organism type
river - gravel / sand - < 0.05 limnetic freshwater
tidal river (seaward of tidal limit) 10 - 100 sand silt / mud 0.05 - 0.5 limnetic freshwater
estuary head (seaward of salt intrusion limit) 2 - 5 sand / silt silt / mud 0.5 - 5 oligohaline (brackish) oligohaline, freshwater migrants
upper reach 5 - 25 sand /silt / mud silt / mud 5 - 18 mesohaline true estuarine, limit of non-transient migrants
middle reach 5 - 25 sand / silt / mud silt / mud 18 - 25 polyhaline estuarine, euryhaline
lower reach 5 - 10 sand silt / mud 25 – 30 polyhaline estuarine, euryhaline, marine migrants
mouth / sea 5 - 10 sand / gravel / rock - 30-35 euhaline stenohaline, all marine


The challenges of estuarine ecosystems

Fig. 2. Typical mudflat organisms.

Organisms that live in estuaries must be adapted to the specific conditions that occur here. The most constraining condition is the varying salinity. Most organisms are so-called stenohalines: they survive only in an environment with a certain more or less constant salinity, for example in the sea or in rivers. These species cannot survive in an estuarine environment. Species that thrive in an estuarine environment must be able to cope with varying osmotic pressures related to varying salinity; these species are called euryhalines. When the internal osmotic pressure of a cell is lower than the external pressure, the cell will lose fluid and shrink; in the opposite case, the cell will swell. There are two strategies to counter this. One strategy is used by the so-called osmoconformers: they regulate the internal osmotic pressure in such a way that it becomes equal to the external osmotic pressure. Depending on the ambient salinity, they achieve this by producing the required amount of organic osmolytes (molecules such as sugars, amino acids, methylamines) to balance the internal and external osmotic pressure[3]. The other strategy is used by the so-called osmoregulators: they regulate the osmotic pressure by secreting an excess of salt or an excess of water, depending on the salinity of the environment. Because euryhaline species are relatively rare, the estuarine ecosystem is characterized by a relatively limited biodiversity (Fig. 2) but a high abundance of specific estuarine species.

Most marine invertebrates are osmoconformers and generally stenohaline. However, some species are euryhaline, for example barnacles, some polychaetes worms and some copepods. Bivalve mollusks are euryhaline osmoconformers within a limited salinity range, but cope with strong salinity variations by closing their valves. Marine vertebrates are generally osmoregulators. Several crustacean species (e.g. crabs) and fish species (e.g. salmon, flounder) are euryhaline. Euryhaline osmoregulating vegetation species include Spartina and Salicornia. Mangroves, however, are adapted to specific salinity ranges and thus cannot be classified as euryhaline.


Lowland reaches of rivers are characterised by high levels of suspended solids, such as silt, clay and organic detritus, inducing high turbidity. In the fresh water – sea water transition zone these particles become effectively ‘trapped’ due to flocculation and converging suspended sediment fluxes [4][5]. These convergences result in a so-called estuarine turbidity maximum (ETM). For further explanation, see the articles Estuarine turbidity maximum and Dynamics of mud transport.

Turbidity limits the depth of the photic zone (light penetration zone), thus limiting the photosynthesis and primary production. Much of the primary production in estuaries occurs on the seaward side of the ETM (Swart, 2007[6]). Macrophyte vegetation and benthic algae are often limited to the periodically exposed (intertidal) part of the estuary, while the growth of phytoplankton is restricted to a thin uppermost layer of the water column (Cloern, 1997[7].) As a result, estuaries are heterotrophic systems, where more energy is consumed than produced. Most estuarine species are detritivores, meeting their energy intake requirements from organic matter contained either in the sediment (deposit feeders), in suspension (filter feeders) or both. Benthic invertebrates play a major role in energy cycling in estuaries (Wilson & Parkes, 1998[8]).

Fig. 3. Mudflat in the Deben estuary (UK).

Particulate matter eventually settles in the estuarine mid reaches where the substratum can contain a high mud fraction, especially in the intertidal areas adjacent to the main tidal channels (Fig. 3). These mudflats are less suitable for organisms with an affinity for hard substratum but provide an important habitat for a range of burrowing invertebrate species, of which bivalve mollusks and polychaete worms are usually dominant in terms of numbers and biomass. These in turn provide an attractive dish for higher trophic consumers.

Sediment deposition gradually increases the height of the intertidal areas until submergence only occurs at the highest tides. Accretion is stimulated by the development of vegetation on the landward zones which are most sheltered from currents and waves. These high vegetated zones of the intertidal area, the so-called saltmarshes, form an important estuarine habitat. This habitat is described in the article Salt marshes.


Industrialised and urbanised river catchments and estuaries receive anthropogenic input from various sources (both point sources and diffuse sources) and often carry a range of contaminants. These pollutants bind to fine sediments and therefore can accumulate in sheltered, low-energy areas such as intertidal mudflats in enclosed bays or estuaries. The estuarine benthic fauna of these zones can thus be subject to high contaminant levels (Kausch and Michaelis, 1996[9]). For further details, see: Coastal and marine sediments.


Biodiversity of estuarine ecosystems

Fig. 4. Species richness in estuaries follows a long established pattern along the salinity gradient described by Remane (1934[10])

Truly estuarine species are those that complete their whole life cycle within the transitional waters. Species permanently dwelling there are mostly hardy, stress-tolerant species able to handle salinity fluctuations and high suspended solid levels. Organisms living on the higher parts of the mudflats and on the salt marshes are subject to additional stresses during temporary emersion at low tide, such as exposure to air, dehydration and temperature variations. Not many species can perform well under such conditions. Estuarine ecosystems are thus characterised by relatively low species diversity compared to freshwater or full salinity conditions. Along the estuary, from head to mouth, freshwater species become rare as salinity increases and are gradually replaced by marine organisms in the lower estuarine reaches, with some truly estuarine species found at intermediate salinities. This pattern is reflected by the overall species richness, where the least diverse fauna is found at salinity levels between 5 and 18 ppt. This is reflected in the so-called Remane curve, see Fig. 4.


Apart from the permanent dwellers, estuaries are host to a number of visitors. Some of them have to travel through estuaries on their migratory route, being either anadromous (spawn in freshwater and feed and grow at sea) or catadromous (spawn at sea and feed and grow in freshwater). The absence of many of the marine predators and rich particulate food supply offer attractive spawning and nursery grounds for many species that normally live under full salinity conditions. Even though estuarine ecosystems are usually species-poor, they maintain a high productive throughput of invertebrate fauna. This productivity provides rich feeding opportunities for a range of higher consumers. Some marine predators are well equipped to cope with reduced salinity and frequently enter estuaries in search of food. Ebb tide makes estuarine beds available to terrestrial predators, of which birds take the greatest share. Estuarine sand and mudflats that are periodically exposed to air support high densities of a diverse bird fauna. This value of estuaries has been long recognized and many estuarine sand and mudflats have been designated as Special Protection Areas (SPA) under the EU Birds Directive (79/409/EEC).


Threats

Estuarine ecosystems are usually dominated by stress-tolerant organism, able to withstand a wide range of environmental conditions. However, they also have to face serious threats from anthropogenic activities.

Fig. 5. Historical maps of the Loire estuary channelization. Photo: Groyne field along the northern channel bend (at the right in the maps).


  • Estuaries are preferred locations for human settlement, especially due to the natural connections they offer between inland and overseas destinations. Residential, recreational and industrial developments (such as harbours or ports and marinas) are usually located right at the waterfront with supporting structures (roads, railways) that cut off the connections with the upper shore habitats. Estuaries are often challenged by land development; land reclamation is particularly detrimental in this respect as it results in a permanent loss of habitat.


  • Making estuaries with shifting shoals and channels navigable has long been beyond human capabilities. This changed at the end of the 19th century, when large-scale dredging technology became available. Without exception, all estuaries on which harbors were located, have been channeled in the 20th century (Fig. 5). A deep navigation channel was dredged, often deviating from the natural tidal channel. The channel banks were set in stone and groynes were constructed to concentrate the flow and to suppress the natural tendency of tidal channels to shift and meander. Channel deepening allowed the tide to propagate further up the river (with especially a strong lowering in the low water level) and both salt intrusion and turbidity increased, sometimes to a very large extent. Fixing the channel banks has led to strong sedimentation of the remaining intertidal areas, causing the mud flat and salt marsh levels to rise. The raised salt marshes have often been embanked and turned into agricultural land or have become terrestrial habitats. The dynamics of migrating tidal channels, the natural agent of habitat rejuvenation and new marsh development, has therefore been thwarted and replaced by a development towards aging landscapes with significant loss of biodiversity.


  • Rivers discharging into their estuaries carry various constituents depending on the landuse of the drainage area (catchment). This means that various contaminants introduced at any point in the catchment ultimately end up in the estuary. Although estuarine organisms are typically hardy, these pollutants and excess nutrients impede their overall performance (including growth and reproduction). Pollution from densely populated or heavily industralised catchments has detrimental effects on life in estuaries. See: Eutrophication in coastal environments, Coastal pollution and impacts.


  • Climate change impacts of concern for estuaries are the overall temperature rise and elevation of the sea level. The former is likely to induce latitudinal migration with more warm-water species being increasingly established and possibly outcompeting the native species in a long run. Sea level rise would result in a shift of the salinity zonation landwards (see also Effects of global climate change on European marine biodiversity). However, coastal areas, including estuaries, are usually heavily populated and developed. In such developed areas the vertical shift of salinity zonation will be hindered by flood defence structures and river and shore embankments resulting in estuarine squeeze. Because many estuaries are situated in lowland, a substantial rise of the high water levels will have great consequences for the safety of the people living in these low-lying areas and for their properties and infrastructure. See the article Coastal cities and sea level rise.


Related articles

Morphology of estuaries
Estuarine turbidity maximum
Salt marshes
Mangroves
Blue carbon sequestration
Tidal asymmetry and tidal basin morphodynamics
Coastal and marine sediments
Estuarine morphological modelling
Osmosis
Other article in the category Estuaries and tidal rivers


References

  1. Carriker, M. R. 1967. Ecology of estuarine benthic invertebrates: a perspective, p. 442-487. In G. H. Lauff (ed.), Estuaries, American Association for the Advancement of Science. Washington.
  2. McLusky, D. S. 1989. The Estuarine Ecosystem, 2nd Ed. Chapman & Hall, London.
  3. Yancey, P.H. 2005. Organic osmolytes as compatible, metabolic and counteracting cytoprotectants in high osmolarity and other stresses. J. Exp. Biol. 208: 2819–2830
  4. Kranck, K. (1981) Particulate matter grain-size characteristics and flocculation in a partially mixed estuary. Sedimentology 28: 107 – 114.
  5. Robinson M. C., Morris, K. P. & Dyer, K. R. (1999). Deriving fluxes of suspended particulate matter in the Humber Estuary, UK, using airborne remote sensing. Marine Pollution Bulletin, 37: 155 -163
  6. Swart, de H. E., Schuttelaars, H. M. & Talke, S. A. (2007). A simple model for phytoplankton growth in turbid estuaries. Geophysical Research Abstracts 9, 04190.
  7. Cloern, J.E. (1997). Turbidity as a control on phytoplankton biomass and productivity in estuaries. Continental Shelf Research 7: 1367-1381.
  8. Wilson, J.G. & Parkes, A. (1998). Network analysis of the energy flow through the Dublin Bay ecosystem. Biology And Environment: Proceedings Of The Royal Irish Academy 98B (3): 179–190.
  9. Kausch, H. and Michaelis, W., eds. 1996. Suspended particulate matter in rivers and estuaries - Proceedings of an International Symposium held at Reinbek near Hamburg, Germany. Advances in Limnology, 47.
  10. Remane, A. (1934). Die Brackwasserfauna. Verzeichnis der Veröffentlichungen Goldsteins, 36: 34–74. .


The main author of this article is Penk, Marcin
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Citation: Penk, Marcin (2021): Estuarine ecosystems. Available from http://www.coastalwiki.org/wiki/Estuarine_ecosystems [accessed on 28-03-2024]