Wikipedia

Filter feeder

Filter feedersareaquatic animalsthat acquirenutrientsbyfeedingonorganic matters,foodparticlesor smallerorganisms(bacteria,microalgaeandzooplanktons)suspendedin water, typically by having the water pass over or through a specializedfilteringorganthat sieves out and/or traps solids. Filter feeders can play an important role in condensingbiomassand removingexcess nutrients(such asnitrogenandphosphate) from the localwaterbody,and are therefore considered water-cleaningecosystem engineers.They are also important inbioaccumulationand, as a result, asindicator organisms.

Krill feeding in a highphytoplanktonconcentration (slowed by a factor of 12)

Filter feeders can besessile,planktonic,nektonicor evenneustonic(in the case of thebuoy barnacle) depending on thespeciesand thenichesthey haveevolvedto occupy. Extant species that rely on suchmethod of feedingencompass numerousphyla,includingporiferans(sponges),cnidarians(jellyfish,sea pensandcorals),arthropods(krill,mysidsandbarnacles),molluscs(bivalves,such asclams,scallopsandoysters),echinoderms(sea lilies) andchordates(lancelets,sea squirtsandsalps,as well as manymarinevertebratessuch as most species offorage fish,American paddlefish,silverandbighead carps,baleen whales,manta rayand three species ofsharks—thewhale shark,basking sharkandmegamouth shark). Somewater birdssuch asflamingosand certainduckspecies, though predominantly terrestrial, are also filter feeders whenforaging.

Fish

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See also:Forage fish

Mostforage fishare filter feeders. For example, theAtlantic menhaden,a type ofherring,lives onplanktoncaught in midwater. Adult menhaden can filter up to four gallons of water a minute and play an important role in clarifying ocean water. They are also a natural check to the deadlyred tide.[1]

In addition to these bony fish, four types ofcartilaginous fishesare also filter feeders. Thewhale sharksucks in a mouthful of water, closes its mouth and expels the water through itsgills.During the slight delay between closing the mouth and opening the gill flaps, plankton is trapped against thedermal denticleswhich line its gill plates andpharynx.This fine sieve-like apparatus, which is a unique modification of the gill rakers, prevents the passage of anything but fluid out through the gills (anything above 2 to 3 mm in diameter is trapped). Any material caught in the filter between the gill bars is swallowed. Whale sharks have been observed "coughing" and it is presumed that this is a method of clearing a build up of food particles in the gill rakers.[2][3][4]Themegamouth sharkhas luminous organs calledphotophoresaround its mouth. It is believed they may exist to lure plankton or small fish into its mouth.[5]Thebasking sharkis a passive filter feeder, filteringzooplankton,small fish, andinvertebratesfrom up to 2,000 tons of water per hour.[6]Unlike the megamouth and whale sharks, the basking shark does not appear to actively seek its quarry; but it does possess largeolfactory bulbsthat may guide it in the right direction. Unlike the other large filter feeders, it relies only on the water that is pushed through the gills by swimming; the megamouth shark and whale shark can suck or pump water through their gills.[6]Manta rayscan time their arrival at the spawning of large shoals of fish and feed on the free-floating eggs and sperm. This stratagem is also employed by whale sharks.[7]

Arthropods

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Filter basket of amysid

Like all arthropods, crustaceans areecdysozoans,a clade withoutcilia.Cilia play an important role for many filter feeding animals, but because crustaceans don't have them, they need to use modified extremities for filter feeding instead.[8]Mysidaceanslive close to shore and hover above the sea floor, constantly collecting particles with their filter basket. They are an important food source forherring,cod,flounder,andstriped bass.Mysids have a high resistance to toxins in polluted areas, and may contribute to high toxin levels in their predators.[citation needed]Antarctic krillmanages to directly utilize the minutephytoplanktoncells, which no other higher animal of krill size can do. This is accomplished through filter feeding, using the krill's developed front legs, providing for a very efficient filtering apparatus:[9]the sixthoracopodsform a very effective "feeding basket" used to collect phytoplankton from the open water. In the animation at the top of this page, the krill is hovering at a 55° angle on the spot. In lower food concentrations, the feeding basket is pushed through the water for over half a meter in an opened position, and then the algae are combed to the mouth opening with special setae on the inner side of the thoracopods. Porcelain crabshave feeding appendages covered with setae to filter food particles from the flowing water.[10] Most species ofbarnaclesare filter feeders, using their highly modified legs to sift plankton from the water.[11]

Also some insects with aquatic larvae or nymphs are filter feeders during their aquatic stage. Such as some species ofmayflynymphs,[12]mosquitolarvae,[13]andblack flylarvae.[14]Instead of using modified limbs or mouthparts, somecaddisflylarvae produce nets of silk used for filter feeding.[15]

Baleen whales

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Mouth plates of a baleen whale

Thebaleen whales(Mysticeti), one of two suborders of theCetacea(whales, dolphins, and porpoises), are characterized by havingbaleenplates for filtering food from water, rather than teeth. This distinguishes them from the other suborder of cetaceans, thetoothed whales(Odontoceti). The suborder contains four families and fourteen species. Baleen whales typically seek out a concentration of zooplankton, swim through it, either open-mouthed or gulping, and filter the prey from the water using their baleens. A baleen is a row of a large number ofkeratinplates attached to the upper jaw with a composition similar to those in human hair or fingernails. These plates are triangular in section with the largest, inward-facing side bearing fine hairs forming a filtering mat.[16]Right whalesare slow swimmers with large heads and mouths. Their baleen plates are narrow and very long — up to 4 m (13 ft) inbowheads— and accommodated inside the enlarged lower lip which fits onto the bowed upper jaw. As the right whale swims, a front gap between the two rows of baleen plates lets the water in together with the prey, while the baleens filter out the water.[16]Rorqualssuch as theblue whale,in contrast, have smaller heads, are fast swimmers with short and broad baleen plates. To catch prey, they widely open their lower jaw — almost 90° — swim through a swarm gulping, while lowering their tongue so that the head's ventral grooves expand and vastly increase the amount of water taken in.[16]Baleen whales typically eatkrillin polar or subpolar waters during summers, but can also take schooling fish, especially in the Northern Hemisphere. All baleen whales except thegray whalefeed near the water surface, rarely diving deeper than 100 m (330 ft) or for extended periods. Gray whales live in shallow waters feeding primarily on bottom-living organisms such asamphipods.[16]

Bivalves

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External image
Movie clip of siphon feeding

Bivalvesare aquaticmolluscswhich havetwo-part shells.Typically both shells (or valves) aresymmetricalalong the hinge line. The class has 30,000species,includingscallops,clams,oystersandmussels.Most bivalves are filter feeders (although some have taken up scavenging and predation), extracting organic matter from the sea in which they live.Nephridia,the shellfish version ofkidneys,remove the waste material. Buried bivalves feed by extending a siphon to the surface. For example,oystersdraw water in over their gills through the beating ofcilia.Suspended food (phytoplankton,zooplankton,algaeand other water-borne nutrients and particles) are trapped in the mucus of a gill, and from there are transported to the mouth, where they are eaten, digested and expelled as feces orpseudofeces.Each oyster filters up to five litres of water per hour. Scientists believe that theChesapeake Bay's once-flourishing oyster population historically filtered the estuary's entire water volume of excess nutrients every three or four days. Today that process would take almost a year,[17]and sediment, nutrients, and algae can cause problems in local waters. Oysters filter these pollutants,[18]and either eat them or shape them into small packets that are deposited on the bottom where they are harmless.

Bivalve shellfish recycle nutrients that enter waterways from human and agricultural sources.Nutrient bioextractionis "an environmental management strategy by which nutrients are removed from an aquatic ecosystem through the harvest of enhanced biological production, including the aquaculture of suspension-feeding shellfish or algae".[19]Nutrient removal by shellfish, which are then harvested from the system, has the potential to help address environmental issues including excess inputs of nutrients (eutrophication), low dissolved oxygen, reduced light availability and impacts on eelgrass, harmful algal blooms, and increases in incidence ofparalytic shellfish poisoning(PSP). For example, the average harvested mussel contains: 0.8–1.2% nitrogen and 0.06–0.08% phosphorus[20]Removal of enhanced biomass can not only combat eutrophication and also support the local economy by providing product for animal feed or compost. In Sweden, environmental agencies utilize mussel farming as a management tool in improving water quality conditions, wheremussel bioextractionefforts have been evaluated and shown to be a highly effective source of fertilizer and animal feed[21]In the U.S., researchers are investigating potential to model the use of shellfish and seaweed for nutrient mitigation in certain areas of Long Island Sound.[22]

Bivalves are also largely used asbioindicatorsto monitor the health of an aquatic environment, either fresh- or seawater. Their population status or structure, physiology, behaviour,[23]or their content of certain elements or compounds can reveal the contamination status of any aquatic ecosystem. They are useful as they are sessile, which means they are closely representative of the environment where they are sampled or placed (caging), and they breathe water all the time, exposing their gills and internal tissues:bioaccumulation.One of the most famous projects in that field is theMussel Watch Programmein America.

Sponges

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Tube sponges attracting smallreef fish

Sponges have no truecirculatory system;instead, they create a water current which is used for circulation. Dissolved gases are brought to cells and enter the cells via simplediffusion.Metabolic wastesare also transferred to the water through diffusion. Sponges pump remarkable amounts of water.Leuconia,for example, is a small leuconoid sponge about 10 cm tall and 1 cm in diameter. It is estimated that water enters through more than 80,000 incurrent canals at a speed of 6 cm per minute. However, becauseLeuconiahas more than 2 million flagellated chambers whose combined diameter is much greater than that of the canals, water flow through chambers slows to 3.6 cm per hour.[24]Such a flow rate allows easy food capture by the collar cells. Water is expelled through a singleosculumat a velocity of about 8.5 cm/second: a jet force capable of carrying waste products some distance away from the sponge.

Cnidarians

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Themoon jellyfishhas a grid of fibres which are slowly pulled through the water. The motion is so slow thatcopepodscannot sense it and do not react with anescape response.[citation needed]

  • An undulating liveAureliain theBaltic Seashowing the grid in action.
  • Higher magnification showing a prey item, probably acopepod.
  • The prey is then drawn to the body by contracting the fibres in a corkscrew fashion (image taken with anecoSCOPE).

Other filter-feeding cnidarians includesea pens,sea fans,plumose anemones,andXenia.[citation needed]

Tunicates

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Tunicates take water in through a siphon and then expel filtered water through another siphon

Tunicates,such asascidians,salpsandsea squirts,arechordateswhich form a sister group to thevertebrates.Nearly all tunicates aresuspension feeders,capturingplanktonicparticles by filtering sea water through their bodies. Water is drawn into the body through the inhalantbuccalsiphon by the action ofcilialining the gill slits. The filtered water is then expelled through a separate exhalant siphon. To obtain enough food, a typical tunicate needs to process about one body-volume of water per second.[25]

Birds

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Thearcuatebill of thislesser flamingois well adapted to bottom scooping

Flamingosfilter-feed onbrine shrimp.Their oddly shaped beaks are specially adapted to separate mud and silt from the food they eat, and are uniquely used upside-down. The filtering of food items is assisted by hairy structures calledlamellaewhich line themandibles,and the large rough-surfaced tongue.[26]

Prionsare specialised petrels with filter-feeding habits. Their name comes from their saw-like jaw edges, used to scope out small planktionic animals.[27]

The extinctswanAnnakacygnais speculated to be a filter-feeder due to its bill proportions being similar to those ofshoveler ducks.It is unique in being a large, flightless marine animal, unlike the smaller still volant flamingos and prions.

Pterosaurs

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Traditionally,Ctenochasmatoideaas a group has been listed as filter-feeders, due to their long, multiple slender teeth, clearly well adapted to trap prey. However, onlyPterodaustroshowcases a proper pumping mechanism, having up-turned jaws and powerful jaw and tongue musculature. Other ctenochasmatoids lack these, and are now instead thought to have beenspoonbill-like catchers, using their specialised teeth simply to offer a larger surface area. Tellingly, these teeth, while small and numerous, are comparatively unspecialised to the baleen-like teeth ofPterodaustro.[28]

Boreopteridsare thought to have relied on a kind of rudimentary filter feeding, using their long, slender teeth to trap small fish, though probably lacking the pumping mechanism ofPterodaustro.In essence, their foraging mechanism was similar to that of modern youngPlatanista"dolphins".[28][29]

Marine reptiles

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Filter feeding habits are conspicuously rare amongMesozoicmarine reptiles,the main filter feeding niche being seemingly instead occupied bypachycormidfish. However, some sauropsids have been suggested to have engaged in filter feeding.Henoduswas a placodont with unique baleen-like denticles and features of the hyoid and jaw musculature comparable to those of flamingos. Combined with its lacustrine environment, it might have occupied a similar ecological niche.[30][31]In particular, it was probably aherbivore,filtering outalgaeand other small-sizedflorafrom the substrates.[32] Stomatosuchidaeis a family of freshwatercrocodylomorphswith rorqual-like jaws and minuscule teeth, and the unrelatedCenozoicMourasuchusshares similar adaptations. Hupehsuchiais a lineage of bizarreTriassicreptiles adapted for suspension feeding.[33] Someplesiosaursmight have had filter-feeding habits.[34]

See also

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Citations

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  1. ^H. Bruce Franklin (March 2006)."Net Losses: Declaring War on the Menhaden".Mother Jones.Retrieved27 February2009.Extensive article on the role of menhaden in the ecosystem and possible results of overfishing.
  2. ^Ed. Rainer Froese and Daniel Pauly."Rhincodon typus".FishBase.Retrieved17 September2006.
  3. ^Martin, R. Aidan."Elasmo Research".ReefQuest.Retrieved17 September2006.
  4. ^"Whale shark".Ichthyology at the Florida Museum of Natural History. Archived fromthe originalon 5 September 2006.Retrieved17 September2006.
  5. ^Bird, Christopher (28 October 2014)."Glow in the Dark Sharks".University of Southampton.Retrieved11 June2018.
  6. ^abC. Knickle; L. Billingsley; K. DiVittorio."Biological Profiles basking shark".Florida Museum of Natural History. Archived fromthe originalon 21 August 2006.Retrieved11 June2018.
  7. ^Hall, Danielle."The Massive Filter Feeding Shark You Ought to Know | Smithsonian Ocean".ocean.si.edu.Retrieved30 August2022.
  8. ^Neuronal coordination of motile cilia in locomotion and feeding
  9. ^Kils, U.:Swimming and feeding of Antarctic Krill,Euphausia superba- some outstanding energetics and dynamics—some unique morphological details.InBerichte zur Polarforschung,Alfred Wegener Institute for Polar and Marine Research,Special Issue 4 (1983): "On the biology of KrillEuphausia superba",Proceedings of the Seminar and Report of Krill Ecology Group, Editor S. B. Schnack, 130-155 and title page image.
  10. ^Valdivia, Nelson; Stotz, Wolfgang (2006)."Feeding Behavior of the Porcellanid Crab Allopetrolisthes Spinifrons, Symbiont of the Sea Anemone Phymactis Papillosa".Journal of Crustacean Biology.26(3): 308–315.doi:10.1651/C-2607.1.
  11. ^"Acorn Barnacles".Field Studies Council. 2008.Retrieved11 June2018.
  12. ^Sroka, Pavel; Staniczek, Arnold H. (2022)."Evolution of filter-feeding in aquatic insects dates back to the Middle Triassic: New evidence from stem-group mayflies (Insecta, Ephemerida) from Grès à Voltzia, Vosges, France".Papers in Palaeontology.8(4).Bibcode:2022PPal....8E1456S.doi:10.1002/spp2.1456.
  13. ^Roberts, Derek (2014)."Mosquito Larvae Change Their Feeding Behavior in Response to Kairomones From Some Predators".Journal of Medical Entomology.51(2): 368–374.doi:10.1603/ME13129.PMID24724285.
  14. ^Kurtak, Daniel C. (1978)."Efficiency of filter feeding of black fly larvae (Diptera: Simuliidae)".Canadian Journal of Zoology.56(7): 1608–1623.doi:10.1139/z78-222.
  15. ^Mason, Richard (June 2020).The zoogeomorphology of case-building caddisfly larvae(PDF)(Thesis).
  16. ^abcdBannister, John L. (2008). "Baleen Whales (Mysticetes)". In Perrin, William F.; Würsig, Bernd; Thewissen, J. G. M. (eds.).Encyclopedia of Marine Mammals.Academic Press. pp.80–89.ISBN978-0-12-373553-9.
  17. ^"Oyster Reefs: Ecological importance".US National Oceanic and Atmospheric Administration.Retrieved11 June2018.
  18. ^The comparative roles of suspension-feeders in ecosystems. Springer. Dordrecht, 359 p.
  19. ^NOAA. "Nutrient Bioextraction Overview".Long Island Sound Study.
  20. ^Stadmark and Conley. 2011. Mussel farming as a nutrient reduction measure in the Baltic Sea: consideration of nutrient biogeochemical cycles. Marine Pollution Bull. 62(7):1385-8
  21. ^Lindahl O, Hernroth R., Kollberg S., Loo L.-O, Olrog L., Rehnstam-Holm A.-S., Svensson J., Svensson S., Syversen U. (2005). "Improving marine water quality by mussel farming: A profitable solution for Swedish society".Ambio.34(2): 131–138.Bibcode:2005Ambio..34..131L.doi:10.1579/0044-7447-34.2.131.PMID15865310.S2CID25371433.{{cite journal}}:CS1 maint: multiple names: authors list (link)
  22. ^Miller and Wands."Applying the System Wide Eutrophication Model (SWEM) for a Preliminary Quantitative Evaluation of Biomass Harvesting as a Nutrient Control Strategy for Long Island Sound"(PDF).Hydroqual, Inc.
  23. ^"behaviour".Archived fromthe originalon 13 November 2016.Retrieved25 January2014.
  24. ^See Hickman and Roberts (2001) Integrated principles of zoology – 11th ed., p. 247
  25. ^Ruppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004).Invertebrate Zoology, 7th edition.Cengage Learning. pp. 940–956.ISBN978-81-315-0104-7.{{cite book}}:CS1 maint: multiple names: authors list (link)
  26. ^Carnaby, Trevor (2010) [2008].Beat about the Bush: Birds.Jacana. p. 456.ISBN978-1-77009-241-9.
  27. ^Gotch, A. F. (1995) [1979]. "Albatrosses, Fulmars, Shearwaters, and Petrels". Latin Names Explained A Guide to the Scientific Classifications of Reptiles, Birds & Mammals. New York, NY: Facts on File. pp. 191–192. ISBN 0-8160-3377-3.
  28. ^abWilton, Mark P. (2013).Pterosaurs: Natural History, Evolution, Anatomy.Princeton University Press.ISBN978-0691150611.
  29. ^Pilleri G., Marcuzzi G., Pilleri O. (1982). "Speciation in the Platanistoidea, systematic, zoogeographical and ecological observations on recent species".Investigations on Cetacea.14:15–46.{{cite journal}}:CS1 maint: multiple names: authors list (link)
  30. ^Rieppel, O. (2002). Feeding mechanisms in Triassic stem-group sauropterygians: the anatomy of a successful invasion of Mesozoic seas Zoological Journal of the Linnean Society, 135, 33-63
  31. ^Naish D (2004). "Fossils explained 48. Placodonts".Geology Today.20(4): 153–158.Bibcode:2004GeolT..20..153N.doi:10.1111/j.1365-2451.2004.00470.x.S2CID128475420.
  32. ^Chun, Li; Rieppel, Olivier; Long, Cheng; Fraser, Nicholas C. (May 2016)."The earliest herbivorous marine reptile and its remarkable jaw apparatus".Science Advances.2(5): e1501659.Bibcode:2016SciA....2E1659C.doi:10.1126/sciadv.1501659.PMC4928886.PMID27386529.
  33. ^Sanderson S. L., Wassersug R. (1990). "Suspension-feeding vertebrates".Scientific American.262(3): 96–101.Bibcode:1990SciAm.262c..96S.doi:10.1038/scientificamerican0390-96.
  34. ^"Plesiosaur Machinations XI: Imitation Crab Meat Conveyor Belt and the Filter Feeding Plesiosaur".25 July 2015.Retrieved11 June2018.
  35. ^Gregorič, Matjaž; Kutnjak, Denis; Bačnik, Katarina; Gostinčar, Cene; Pecman, Anja; Ravnikar, Maja; Kuntner, Matjaž (16 May 2022). "Spider webs as eDNA samplers: Biodiversity assessment across the tree of life".Molecular Ecology Resources.22(7). Wiley: 2534–2545.doi:10.1111/1755-0998.13629.ISSN1755-098X.PMID35510791.S2CID248527088.

General and cited references

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