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Filter feeders are a sub-group of suspension feeding
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feeding are clams, krill,
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throug ...
s,
baleen whale Baleen whales ( systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea ( whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in th ...
s, and many
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...
(including some
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s). Some birds, such as flamingos and certain species of duck, are also filter feeders. Filter feeders can play an important role in clarifying water, and are therefore considered ecosystem engineers. They are also important in
bioaccumulation Bioaccumulation is the gradual accumulation of substances, such as pesticides or other chemicals, in an organism. Bioaccumulation occurs when an organism absorbs a substance at a rate faster than that at which the substance is lost or eliminated ...
and, as a result, as indicator organisms.


Fish

Most
forage fish Forage fish, also called prey fish or bait fish, are small pelagic fish which are preyed on by larger predators for food. Predators include other larger fish, seabirds and marine mammals. Typical ocean forage fish feed near the base of the ...
are filter feeders. For example, the Atlantic menhaden, a type of
herring Herring are forage fish, mostly belonging to the family of Clupeidae. Herring often move in large schools around fishing banks and near the coast, found particularly in shallow, temperate waters of the North Pacific and North Atlantic Ocea ...
, lives on
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cr ...
caught 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 deadly
red tide A harmful algal bloom (HAB) (or excessive algae growth) is an algal bloom that causes negative impacts to other organisms by production of natural phycotoxin, algae-produced toxins, mechanical damage to other organisms, or by other means. HABs are ...
. Extensive article on the role of menhaden in the ecosystem and possible results of overfishing. In addition to these bony fish, four types of
cartilaginous fishes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue ...
are also filter feeders. The whale shark sucks in a mouthful of water, closes its mouth and expels the water through its gills. During the slight delay between closing the mouth and opening the gill flaps, plankton is trapped against the dermal denticles which line its gill plates and
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its ...
. 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. The megamouth shark has luminous organs called photophores around its mouth. It is believed they may exist to lure plankton or small fish into its mouth. The basking shark is a passive filter feeder, filtering
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, small fish, and
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s from up to 2,000 tons of water per hour. Unlike the megamouth and whale sharks, the basking shark does not appear to actively seek its quarry; but it does possess large olfactory bulbs that 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. Manta rays can 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.


Crustaceans

Mysidacea are small crustaceans that live close to shore and hover above the sea floor, constantly collecting particles with their filter basket. They are an important food source for
herring Herring are forage fish, mostly belonging to the family of Clupeidae. Herring often move in large schools around fishing banks and near the coast, found particularly in shallow, temperate waters of the North Pacific and North Atlantic Ocea ...
, cod, flounder, and striped bass. Mysids have a high resistance to toxins in polluted areas, and may contribute to high toxin levels in their predators. Antarctic krill manages to directly utilize the minute
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. P ...
cells, 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: the six thoracopods form 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 crabs have feeding appendages covered with setae to filter food particles from the flowing water. Most species of barnacles are filter feeders, using their highly modified legs to sift plankton from the water.


Baleen whales

The
baleen whale Baleen whales ( systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea ( whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in th ...
s (Mysticeti), one of two suborders of the Cetacea (whales, dolphins, and porpoises), are characterized by having baleen plates for filtering food from water, rather than teeth. This distinguishes them from the other suborder of cetaceans, the toothed 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 of keratin plates 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. Right whales are slow swimmers with large heads and mouths. Their baleen plates are narrow and very long — up to in bowheads — 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. Rorquals such as the
blue whale The blue whale (''Balaenoptera musculus'') is a marine mammal and a baleen whale. Reaching a maximum confirmed length of and weighing up to , it is the largest animal known to have ever existed. The blue whale's long and slender body can b ...
, 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. Baleen whales typically eat krill in polar or subpolar waters during summers, but can also take schooling fish, especially in the Northern Hemisphere. All baleen whales except the
gray whale The gray whale (''Eschrichtius robustus''), also known as the grey whale,Britannica Micro.: v. IV, p. 693. gray back whale, Pacific gray whale, Korean gray whale, or California gray whale, is a baleen whale that migrates between feeding and bre ...
feed near the water surface, rarely diving deeper than or for extended periods. Gray whales live in shallow waters feeding primarily on bottom-living organisms such as amphipods.


Bivalves

Bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biv ...
s are aquatic molluscs which have two-part shells. Typically both shells (or valves) are symmetrical along the hinge line. The class has 30,000
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
, including scallops, clams,
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not a ...
s and mussels. 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 of
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; bloo ...
s, remove the waste material. Buried bivalves feed by extending a siphon to the surface. For example,
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not a ...
s draw water in over their gills through the beating of cilia. Suspended food (
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. P ...
,
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
,
algae Algae ( , ; : alga ) are any of a large and diverse group of photosynthetic, eukaryotic organisms. The name is an informal term for a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from ...
and 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 or
pseudofeces Pseudofeces or pseudofaeces are a specialized method of expulsion that filter-feeding bivalve mollusks (and filter-feeding gastropod mollusks) use in order to get rid of suspended particles such as particles of grit which cannot be used as food, ...
. Each oyster filters up to five litres of water per hour. Scientists believe that the
Chesapeake Bay The Chesapeake Bay ( ) is the largest estuary in the United States. The Bay is located in the Mid-Atlantic (United States), Mid-Atlantic region and is primarily separated from the Atlantic Ocean by the Delmarva Peninsula (including the parts: the ...
'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, and sediment, nutrients, and algae can cause problems in local waters. Oysters filter these pollutants, 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 bioextraction is "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". 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 Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytopla ...
), low dissolved oxygen, reduced light availability and impacts on eelgrass, harmful algal blooms, and increases in incidence of paralytic shellfish poisoning (PSP). For example, the average harvested mussel contains: 0.8–1.2% nitrogen and 0.06–0.08% phosphorus 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, where mussel bioextraction efforts have been evaluated and shown to be a highly effective source of fertilizer and animal feed 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. Bivalves are also largely used as bioindicators to monitor the health of an aquatic environment, either fresh- or seawater. Their population status or structure, physiology, behaviour, 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 Bioaccumulation is the gradual accumulation of substances, such as pesticides or other chemicals, in an organism. Bioaccumulation occurs when an organism absorbs a substance at a rate faster than that at which the substance is lost or eliminated ...
. One of the most famous projects in that field is the Mussel Watch Programme in America.


Sponges

Sponges have no true
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
; instead, they create a water current which is used for circulation. Dissolved gases are brought to cells and enter the cells via simple
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 ...
. Metabolic wastes are also transferred to the water through diffusion. Sponges pump remarkable amounts of water. ''
Leuconia ''Leuconia'' is a genus of calcareous sponges in the Family (biology), family Baeriidae. It was described by English anatomist and zoologist Robert Edmond Grant in 1833.Grant, R. E. (1833). Lectures on Comparative Anatomy and Animal Physiology. ...
'', 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, because ''Leuconia'' has 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. Such a flow rate allows easy food capture by the collar cells. Water is expelled through a single osculum at a velocity of about 8.5 cm/second: a jet force capable of carrying waste products some distance away from the sponge.


Cnidarians

The moon jellyfish has a grid of fibres which are slowly pulled through the water. The motion is so slow that
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic (inhabiting sea waters), some are benthic (living on the ocean floor), a number of species have p ...
s cannot sense it and do not react with an escape response. Image:Aureliaauritakils1.jpg, An undulating live ''Aurelia'' in the
Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden and the North and Central European Plain. The sea stretches from 53°N to 66°N latitude and fr ...
showing the grid in action. Image:Aureliaauritakils2.jpg, Higher magnification showing a prey item, probably a
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic (inhabiting sea waters), some are benthic (living on the ocean floor), a number of species have p ...
. Image:Aureliaauritakils3.jpg, The prey is then drawn to the body by contracting the fibres in a corkscrew fashion (image taken with an
ecoSCOPE The ecoSCOPE is an optical sensor system, deployed from a small remotely operated vehicle ( ROV) or fibre optic cable, to investigate behavior and microdistribution of small organisms in the ocean. Deployment Although an ROV may be very small a ...
).
Other filter-feeding cnidarians include sea pens, sea fans, plumose anemones, and '' Xenia''.


Tunicates

Tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one time ...
s, such as
ascidians Ascidiacea, commonly known as the ascidians, tunicates (in part), and sea squirts (in part), is a polyphyletic class in the subphylum Tunicata of sac-like marine invertebrate filter feeders. Ascidians are characterized by a tough outer "tun ...
, salps and sea squirts, are
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...
s which form a sister group to the
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s. Nearly all tunicates are suspension feeders, capturing
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cr ...
ic particles by filtering sea water through their bodies. Water is drawn into the body through the inhalant buccal siphon by the action of cilia lining 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.


Birds

Flamingos filter-feed on brine 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 called lamellae which line the mandibles, and the large rough-surfaced tongue. Prions are specialised petrels with filter-feeding habits. Their name comes from their saw-like jaw edges, used to scope out small planktionic animals. The extinct swan '' Annakacygna'' is speculated to be a filter-feeder due to its bill proportions being similar to those of
shoveler duck The northern shoveler (; ''Spatula clypeata''), known simply in Britain as the shoveler, is a common and widespread duck. It breeds in northern areas of Europe and across the Palearctic and across most of North America, wintering in southern Euro ...
s. It is unique in being a large, flightless marine animal, unlike the smaller still volant flamingos and prions.


Pterosaurs

Traditionally, Ctenochasmatoidea as a group has been listed as filter-feeders, due to their long, multiple slender teeth, clearly well adapted to trap prey. However, only '' Pterodaustro'' showcases 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 been spoonbill-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 of '' Pterodaustro''.
Boreopterid Boreopteridae (meaning "northern wings") is a group of pterodactyloid pterosaurs from the Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. Classification In 2006, Lü Junchang and colleagues named the clade Boreopteridae for the ...
s are 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 of ''Pterodaustro''. In essence, their foraging mechanism was similar to that of modern young '' Platanista'' " dolphins".


Marine reptiles

Filter feeding habits are conspicuously rare among
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Creta ...
marine reptiles, the main filter feeding niche being seemingly instead occupied by pachycormid fish. However, some sauropsids have been suggested to have engaged in filter feeding. '' Henodus'' was 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. In particular, it was probably a herbivore, filtering out
algae Algae ( , ; : alga ) are any of a large and diverse group of photosynthetic, eukaryotic organisms. The name is an informal term for a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from ...
and other small-sized
flora Flora (: floras or florae) is all the plant life present in a particular region or time, generally the naturally occurring (indigenous (ecology), indigenous) native plant, native plants. The corresponding term for animals is ''fauna'', and for f ...
from the substrates.
Stomatosuchidae Stomatosuchidae is an extinct family of neosuchian crocodylomorphs. It is defined as the most inclusive clade containing '' Stomatosuchus inermis'' but not ''Notosuchus terrestris'', '' Simosuchus clarki'', '' Araripesuchus gomesii'', ''Baurusuch ...
is a family of freshwater crocodylomorphs with rorqual-like jaws and minuscule teeth, and the unrelated Cenozoic '' Mourasuchus'' shares similar adaptations. Hupehsuchia is a lineage of bizarre
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
reptiles adapted for suspension feeding. Some plesiosaurs might have had filter-feeding habits.


See also

* Particle (ecology) * Planktivore *
Predation Predation is a biological interaction In ecology, a biological interaction is the effect that a pair of organisms living together in a community have on each other. They can be either of the same species (intraspecific interactions), or o ...
* Spider webs – aerial biofilters, with analogies to aquatic filter feeding


Notes


References

* *Some aspects of water filtering activity of filter-feeders // Hydrobiologia. 2005. Vol. 542, No. 1. pp. 275–286


External links


Filter feeder of krill

Mussel Watch Programme
{{DEFAULTSORT:Filter Feeder Aquatic ecology