Planktivorous
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A planktivore is an aquatic organism that feeds on
plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
ic food, including
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
and
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
. Planktivorous organisms encompass a range of some of the planet's smallest to largest multicellular animals in both the present day and in the past billion years;
basking shark The basking shark (''Cetorhinus maximus'') is the second-largest living shark and fish, after the whale shark. It is one of three Planktivore, plankton-eating shark species, along with the whale shark and megamouth shark. Typically, basking sh ...
s and
copepod Copepods (; meaning 'oar-feet') are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (living in the water column), some are benthos, benthic (living on the sedimen ...
s are just two examples of giant and microscopic organisms that feed upon plankton. Planktivory can be an important mechanism of top-down control that contributes to
trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to redu ...
s in aquatic and marine systems. There is a tremendous diversity of feeding strategies and behaviors that planktivores utilize to capture prey. Some planktivores utilize tides and currents to migrate between estuaries and coastal waters; other aquatic planktivores reside in lakes or reservoirs where diverse assemblages of plankton are present, or migrate vertically in the water column searching for prey. Planktivore populations can impact the abundance and community composition of planktonic species through their predation pressure, and planktivore migrations facilitate nutrient transport between benthic and pelagic habitats. Planktivores are an important link in marine and freshwater systems that connect primary producers to the rest of the food chain. As climate change causes negative effects throughout the global oceans, planktivores are often directly impacted through changes to food webs and prey availability. Additionally, harmful algal blooms (HABs) can negatively impact many planktivores and can transfer harmful toxins from the phytoplankton, to the planktivores, and along up the food chain. As an important source of revenue for humans through tourism and commercial uses in fisheries, many conservation efforts are going on globally to protect these diverse animals known as planktivores. __TOC__


Plankton and planktivory across taxonomic classes


Phytoplankton: prey

Plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
are defined as any type of organism that is unable to swim actively against currents and are thus transported by the physical forcing of tides and currents in the ocean.
Phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
form the lowest trophic level of marine food webs and thus capture light energy and materials to provide food and energy for hundreds of thousands of types of planktivores. Because they require light and abundant nutrients, phytoplankton are typically found in surface waters where light rays can penetrate water. Nutrients that sustain phytoplankton include nitrate, phosphate, silicate, calcium, and micronutrients like iron; however, not all phytoplankton require all these identified nutrients and thus differences in nutrient availability impact phytoplankton
species composition Relative species abundance is a component of biodiversity and is a measure of how common or rare a species is relative to other species in a defined location or community.Hubbell, S. P. 2001. ''The unified neutral theory of biodiversity and biogeog ...
. This class of microscopic, photosynthetic organisms includes
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s,
coccolithophore Coccolithophores, or coccolithophorids, are single-celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the kingdom ...
s,
protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...
s,
cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
,
dinoflagellate The Dinoflagellates (), also called Dinophytes, are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered protists. Dinoflagellates are mostly marine plankton, but they are also commo ...
s, and other microscopic
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
. Phytoplankton conduct photosynthesis via pigments in their cells; phytoplankton can use
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
as well as other accessory photosynthetic pigments like
fucoxanthin Fucoxanthin is a xanthophyll, with formula C42H58O6. It is found as an accessory pigment in the chloroplasts of brown algae and most other heterokonts, giving them a brown or olive-green color. Fucoxanthin absorbs light primarily in the blue-gree ...
,
chlorophyll c Chlorophyll ''c'' refers to forms of chlorophyll found in certain marine algae, including the photosynthetic Chromista (e.g. diatoms and brown algae) and dinoflagellates. These pigments are characterized by their unusual chemical structure, with ...
, alloxanthin, and
carotenoid Carotenoids () are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, corn, tomatoes, cana ...
s, depending on species. Due to their environmental requirements for light and nutrients, phytoplankton are most commonly found near continental margins, the equator, high-latitudes, and nutrient-rich areas. They also form the foundation of the
biological pump The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven Carbon sequestration, sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sedim ...
, which transports carbon to depth in the ocean.


Zooplankton: predators and prey

Zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
("zoo" meaning "animal") are generally consumers of other organisms for food. Zooplankton may consume either phytoplankton or other zooplankton, making them the smallest class of planktivores. They are common to most marine pelagic environments and act as an important step in the food chain to transfer energy up from primary producers to the rest of the marine food web. Some zooplankton remain planktonic for their entire lives, while others eventually grow large enough to swim against currents. For instance, fish are born as planktonic larvae but once they grow large enough to swim, they are no longer considered plankton. Many taxonomic groups (e.g. fishes, krill, corals, etc.) are zooplankton at some point in their lives. For example, oysters begin as planktonic larvae; during this stage when they are considered zooplankton, they consume phytoplankton. Once they mature to adulthood, oysters continue to consume phytoplankton. The spiny water flea is another example of a planktivorous invertebrate. Some of the largest communities of zooplankton exist in high latitude systems like the eastern Bering Sea; pockets of dense zooplankton abundance also exist in the
California Current The California Current () is a cold water Pacific Ocean ocean current, current that moves southward along the western coast of North America, beginning off southern British Columbia and ending off southern Baja California Sur. It is considered an ...
and the
Gulf of Mexico The Gulf of Mexico () is an oceanic basin and a marginal sea of the Atlantic Ocean, mostly surrounded by the North American continent. It is bounded on the northeast, north, and northwest by the Gulf Coast of the United States; on the southw ...
. Zooplankton are, in turn, common prey items for planktivores; they respond to environmental change very rapidly due to their relatively short life spans, and so scientists can track their dynamics to understand what might be occurring in the larger marine food web and environment. The relative ratios of certain zooplankton in the larger zooplankton community can also indicate an environmental change (e.g.,
eutrophication Eutrophication is a general term describing a process in which nutrients accumulate in a body of water, resulting in an increased growth of organisms that may deplete the oxygen in the water; ie. the process of too many plants growing on the s ...
) that may be significant. For instance, an increase in
rotifer The rotifers (, from Latin 'wheel' and 'bearing'), sometimes called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic Coelom#Pseudocoelomates, pseudocoelomate animals. They were first describ ...
abundance in the Great Lakes has been correlated with abnormally high levels of nutrients (eutrophication).


Vertebrates: predators and prey

Many fishes are planktivorous during all or part of their life cycles, and these planktivorous fish are important to human industry and as prey for other organisms in the environment like seabirds and
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
fishes. Planktivores comprise a large component of tropical ecosystems; in the Indo-Australian Archipelago, one study identified 350 planktivorous fish species in one studied grid cell and found that 27% of all fish species in this region were planktivorous. This global study found that coral reef habitats globally have a disproportionate amount of planktivorous fishes. In other habitats, examples of planktivorous fishes include many types of salmon like the
pink salmon Pink salmon or humpback salmon (''Oncorhynchus gorbuscha'') is a species of euryhaline ray-finned fish in the family Salmonidae. It is the type species of the genus ''Oncorhynchus'' (Pacific salmon), and is the smallest and most abundant of t ...
, sandeels,
sardine Sardine and pilchard are common names for various species of small, oily forage fish in the herring suborder Clupeoidei. The term "sardine" was first used in English during the early 15th century; a somewhat dubious etymology says it com ...
s, and silvery lightfish. In ancient systems (read more below), the '' Titanichthys'' was an early massive
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
pelagic planktivore, with a lifestyle similar to that of the modern basking,
whale Whales are a widely distributed and diverse group of fully Aquatic animal, aquatic placental mammal, placental marine mammals. As an informal and Colloquialism, colloquial grouping, they correspond to large members of the infraorder Cetacea ...
, and megamouth
sharks Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
, all of whom are also planktivores. Sea birds can also be planktivores;
least auklet The least auklet (''Aethia pusilla'') is a seabird and the smallest species of auk. It is the most abundant seabird in North America, and one of the most abundant in the world, with a population of around nine million birds. They breed on the isl ...
s, crested auklets,
storm petrel Storm petrel or stormy petrel may refer to one of two bird family (biology), families, both in the order Procellariiformes, once treated as the same family. The two families are: * Northern storm petrels (''Hydrobatidae'') are found in the No ...
s, ancient auklets,
phalarope A phalarope is any of three living species of slender-necked shorebirds in the genus ''Phalaropus'' of the bird family Scolopacidae. Phalaropes are close relatives of the shanks and tattlers, the '' Actitis'' and Terek sandpipers, and also ...
s, and many
penguin Penguins are a group of aquatic flightless birds from the family Spheniscidae () of the order Sphenisciformes (). They live almost exclusively in the Southern Hemisphere. Only one species, the Galápagos penguin, is equatorial, with a sm ...
s are all examples of avian planktivores. Planktivorous seabirds can be indicators of ecosystem status because their dynamics often reflect processes affecting many trophic levels, like the consequences of climate change.
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 ever to have existed. The blue whale's long and slender body can ...
s and
bowhead whale The bowhead whale (''Balaena mysticetus''), sometimes called the Greenland right whale, Arctic whale, and polar whale, is a species of baleen whale belonging to the family Balaenidae and is the only living representative of the genus '' Balaena' ...
s as well as some seals like the
crabeater seal The crabeater seal (''Lobodon carcinophaga''), also known as the krill-eater seal, is a true seal with a circumpolar distribution around the coast of Antarctica. They are the only member of the genus ''Lobodon''. They are medium- to large-sized ( ...
(''Lobodon carcinophagus'') are also planktivorous. Blue whales were recently found to consume a vast amount more plankton than was previously understood, representing an important element of the ocean biogeochemical cycle.


Feeding strategies

As previously mentioned, some plankton communities are well-studied and respond to environmental change very rapidly; understanding unusual plankton dynamics can elucidate potential consequences to planktivorous species and the larger marine food chain. One well-studied planktivore species is the gizzard shad (''Dorosoma cepedianum'') which has a voracious appetite for various forms of plankton across its life cycle. Planktivores can be either obligate planktivores, meaning they can only feed on plankton, or
facultative Facultative means "optional" or "discretionary" (antonym ''obligate''), used mainly in biology in phrases such as: * Facultative (FAC), facultative wetland (FACW), or facultative upland (FACU): wetland indicator statuses for plants * Facultative ...
planktivores, which take plankton when available but eat other types of food as well. In the case of the gizzard shad, they are obligate planktivores when larvae and juveniles, in part due to their very small mouth size; larval gizzard shad are most successful when small zooplankton are present in adequate quantities within their habitat. As they grow, gizzard shad become omnivores, consuming phytoplankton, zooplankton, and larger pieces of nutritious
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
. Adult gizzard shad consume large volumes of zooplankton until it becomes scarce, then start consuming organic debris instead. Larval fishes and
blueback herring The blueback herring, blueback shad, or summer shad (''Alosa aestivalis'') is an anadromous species of herring from the east coast of North America, with a range from Nova Scotia to Florida. Blueback herring form schools and are believed to mi ...
are other well-studied examples of obligate planktivores, whereas fishes like the
ocean sunfish The ocean sunfish (''Mola mola''), also known as the common mola, is one of the largest bony fish in the world. It is the type species of the genus ''Mola'', and one of five extant species in the family Molidae. It was once misidentified as th ...
can alternate between plankton and other food sources (i.e., are facultative planktivores). Facultative planktivores tend to be more opportunistic and live in ecosystems with many types of food sources. Obligate planktivores have fewer options for prey choices; they are typically restricted to marine pelagic ecosystems that have a dominant plankton presence, such as highly productive upwelling regions.


Mechanics of consuming plankton

Planktivores, whether obligate or facultative, obtain food in multiple ways. Particulate feeders eat planktonic items selectively, by identifying plankton and pursuing them in the water column.
Filter feeder Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a s ...
s process large volumes of water internally via different mechanisms, explained below, and strain food items out ''en masse'' or remove food particles from water as it passes by. "Tow-net" filter feeders swim rapidly with mouths open to filter the water, whereas "pumping" filter feeders suck in water via pumping actions. The charismatic
flamingo Flamingos or flamingoes () are a type of wading bird in the family Phoenicopteridae, which is the only extant family in the order Phoenicopteriformes. There are four flamingo species distributed throughout the Americas (including the Caribbe ...
is a pumping filter feeder, using its muscular tongue to pump water along specialized grooves in its bill and pump water back out once plankton have been retrieved. In a different filter feeding process, stationary animals, like corals, use their tentacles to grab plankton particles out of the water column and transfer the particles into their mouth. There are numerous interesting adaptations to remove plankton from the water column. The
phalarope A phalarope is any of three living species of slender-necked shorebirds in the genus ''Phalaropus'' of the bird family Scolopacidae. Phalaropes are close relatives of the shanks and tattlers, the '' Actitis'' and Terek sandpipers, and also ...
s use surface tension feeding to transport particles of prey to their mouth to be swallowed. These birds capture individual particles of plankton held in a droplet of water, suspended in their beaks. They then use a sequence of actions that begin with a quick opening of their beak to increase the surface area of the water droplet encasing prey. The action of stretching out the water droplet ultimately pushes the water and prey to the back of the throat where it can be consumed. These birds also spin around at the water surface, creating their own eddies that draw prey up closer to their beaks. Some species actively hunt plankton: in certain habitats such as the deep open ocean, as mentioned above, the planktivorous
basking shark The basking shark (''Cetorhinus maximus'') is the second-largest living shark and fish, after the whale shark. It is one of three Planktivore, plankton-eating shark species, along with the whale shark and megamouth shark. Typically, basking sh ...
(''Cetorhinus maximus'') track the movements of their prey closely up and down the water column. The
megamouth shark The megamouth shark (''Megachasma pelagios'') is a species of deepwater shark. Rarely seen by Human, humans, it measures around long and is the smallest of the three extant taxon, extant filter-feeding sharks alongside the much larger whale sha ...
(''Megachasma pelagios''), another planktivorous species, adopts a similar feeding strategy that mirrors the movement in the water column of their planktonic prey. Similar to active hunting, some zooplankton, like copepods, are ambush hunters meaning they wait in the water column for prey to come within range and then rapidly attack and consume. Some fishes change their feeding strategy throughout their lives; the
Atlantic menhaden The Atlantic menhaden (''Brevoortia tyrannus'') is a North American species of fish in the herring family, Alosidae. Atlantic menhaden are found in North Atlantic coastal and estuarine waters from Nova Scotia south to northern Florida. They ar ...
(''Brevoortia tyrannus'') is an obligate filter feeder in early life stages, but matures into a particulate feeder. Some fishes, like the northern anchovy (''Engraulis mordax'') can merely modify their feeding behavior depending on the prey or environmental conditions. Some fishes also school together when feeding to help improve contact rates of plankton and simultaneously prevent themselves from predation. Some fishes have gill rakes, an internal filtration structure that assists fishes with capturing plankton prey. The amount of gill rakes can indicate planktivory as well as the typical size of plankton consumed, showing a correlation between gill rake structure and the consumed plankton type.


Nutritional value of plankton

Plankton have highly variable chemical compositions, which impacts their nutritional quality as a food source. Scientists are still understanding how nutritional quality varies with the type of plankton; for example
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
nutritional quality is a controversial topic. The ratios of
phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
and
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
to
carbon Carbon () is a chemical element; it has chemical symbol, symbol C and atomic number 6. It is nonmetallic and tetravalence, tetravalent—meaning that its atoms are able to form up to four covalent bonds due to its valence shell exhibiting 4 ...
within a given plankton determine its nutritional quality. More carbon in an organism relative to these two elements decreases the plankton's nutritional value. Additionally, plankton with higher amounts of
polyunsaturated fatty acid In biochemistry and nutrition, a polyunsaturated fat is a fat that contains a polyunsaturated fatty acid (abbreviated PUFA), which is a subclass of fatty acid characterized by a backbone with two or more carbon–carbon double bonds. Some polyunsa ...
s are typically more energy dense. The nutritional value of plankton does sometimes depend on the nutritional needs of the planktivorous species. For fishes, the nutritional value of plankton is dependent on
docosahexaenoic acid Docosahexaenoic acid (DHA) is an omega−3 fatty acid that is an important component of the human brain, cerebral cortex, skin, and retina. It is given the fatty acid notation 22:6(''n''−3). It can be synthesized from alpha-linolenic acid or ...
, long-chain polyunsaturated fatty acids,
arachidonic acid Arachidonic acid (AA, sometimes ARA) is a polyunsaturated omega−6 fatty acid 20:4(ω−6), or 20:4(5,8,11,14). It is a precursor in the formation of leukotrienes, prostaglandins, and thromboxanes. Together with omega−3 fatty acids an ...
, and
eicosapentaenoic acid Eicosapentaenoic acid (EPA; also icosapentaenoic acid) is an omega−3 fatty acid. In physiological literature, it is given the name 20:5(''n''−3). It also has the trivial name timnodonic acid. In chemical structure, EPA is a carboxylic acid wi ...
with higher concentrations of those chemicals leading to higher nutritional value. However, lipids in plankton prey are not the only required chemical for larval fish; Malzahn et al. found that other nutrients, like phosphorus, were necessary before growth improvements due to lipid concentrations can be realized. Additionally, it has been shown experimentally that the nutritional value of prey is more important than prey abundance for larval fishes. With
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
, plankton may decrease in nutritional quality. Lau et al. discovered that warming conditions and inorganic nutrient depletion in lakes as a result of climate change decreased the nutritional value of plankton communities.


Planktivory across ecological systems


Ancient systems

Planktivory is a common feeding strategy among some of our planet's largest organisms in both the present and the past. Massive
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
organisms like pachycormids have recently been identified as planktivores; some individuals of this group reached lengths upwards of 9 feet. Scientists also recently discovered the fossilized remains of another ancient organism, which they named the "false megamouth" ('' Pseudomegachasma'') shark, and which was likely a filter-feeding planktivore during the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period. This new discovery illuminated planktivory as an example of convergent evolution, whereby distinct lineages evolved to fulfill similar dietary niches. In other words, the false megamouth and its planktivory evolved separate from the ancestors of present-day shark planktivores like the megamouth shark, whale shark, and basking shark, all mentioned above.


Arctic systems

The
Arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
supports productive ecosystems that include many types of planktivorous species. Planktivorous
pink salmon Pink salmon or humpback salmon (''Oncorhynchus gorbuscha'') is a species of euryhaline ray-finned fish in the family Salmonidae. It is the type species of the genus ''Oncorhynchus'' (Pacific salmon), and is the smallest and most abundant of t ...
are common in the Arctic and the
Bering Strait The Bering Strait ( , ; ) is a strait between the Pacific and Arctic oceans, separating the Chukchi Peninsula of the Russian Far East from the Seward Peninsula of Alaska. The present Russia–United States maritime boundary is at 168° 58' ...
and have been suggested to exert significant control on structuring the phytoplankton and zooplankton dynamics in the subarctic North Pacific. Shifts in prey type have also been observed: in northern Arctic regions, salmon are typically piscivorous (consuming other fish) while in the southern Arctic and Bering Strait they are planktivorous.
Capelin The capelin or caplin (''Mallotus villosus'') is a small forage fish of the smelt family found in the North Atlantic, North Pacific and Arctic oceans. In summer, it grazes on dense swarms of plankton at the edge of the ice shelf. Larger capel ...
, ''Mallotus villosus'', are also distributed across much of the Arctic and can exert significant control on zooplankton populations as a result of their planktivorous diet. Capelin have also been seen to exhibit cannibalism on their eggs when other types of preferred plankton sources become less available; alternatively, this behavior may be because increased spawning leads to more eggs in the environment for consumption.
Arctic cod ''Arctogadus glacialis'', known also with ambiguous common names Arctic cod and polar cod, is an Arctic species of fish in the cod Family (biology), family Gadidae, related to the true Gadus, cod (genus ''Gadus''). ''Arctogadus glacialis'' is fo ...
are also important zooplankton consumers and appear to follow aggregations of zooplankton around the region. Planktivorous birds like the fork-tailed storm-petrel and many types of
auk Auks or alcids are birds of the family Alcidae in the order Charadriiformes. The alcid family includes the Uria, murres, guillemots, Aethia, auklets, puffins, and Brachyramphus, murrelets. The family contains 25 extant or recently extinct speci ...
lets are also very common in the Arctic.
Little auk The little auk (Europe) or dovekie (North America) ''Alle alle'' is a small auk, the only member of the genus ''Alle''. ''Alle'' is the Sami name of the long-tailed duck; it is onomatopoeic and imitates the call of the drake duck. Linnaeus was n ...
s are the most common Arctic planktivore species; as they reproduce on land, their planktivory creates an important link between marine and terrestrial nutrient reserves. This link is formed as little auks consume plankton with marine-derived nutrients at sea, then deposit nutrient-rich waste products on land during their reproductive process.


Temperate and sub-arctic systems

In freshwater lake systems, planktivory can be an important forcer of
trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to redu ...
s which can ultimately affect phytoplankton production. Fishes, in these systems, can promote phytoplankton productivity by preying on the zooplankton that control phytoplankton abundances. This is an example of top-down trophic control, where higher trophic organisms like fishes impose control on the abundance of lower trophic organisms, like phytoplankton. Such control on primary production via planktivorous organisms can be important in the functioning of mid-western United States lake systems. Fishes are often the most impactful zooplankton predators, as seen in Newfoundland where three-spine stickleback (''Gasterosteus aculeatus'') predate heavily upon zooplankton. In temperate lakes, the
cyprinid Cyprinidae is a Family (biology), family of freshwater fish commonly called the carp or minnow family, including the carps, the true minnows, and their relatives the barb (fish), barbs and barbel (fish), barbels, among others. Cyprinidae is the ...
and centrarchid fish families are commonly represented among the planktivore community. Planktivores can exert significant competition pressure on organisms in certain lake systems; for instance, in an Idaho lake the introduced planktivorous invertebrate shrimp ''Mysis'' ''relicta'' competes with the native landlocked planktivorous salmon kokanees. Because of the salmon's importance in trophic cycling, the loss of fishes in temperate lake systems could lead to widespread ecological consequences; in this example, such a loss could lead to unchecked predation on plankton by ''Mysis relicta''. Planktivory can also be important in man-made reservoirs. In contrast to deeper and colder natural lakes, reservoirs are warmer, shallower, heavily modified human made systems with different ecosystem dynamics. Gizzard shad, the previously mentioned obligate planktivore, is frequently the most common fish in many reservoir systems. In certain sub-Arctic habitats like deep waters, the planktivorous
basking shark The basking shark (''Cetorhinus maximus'') is the second-largest living shark and fish, after the whale shark. It is one of three Planktivore, plankton-eating shark species, along with the whale shark and megamouth shark. Typically, basking sh ...
tracks the movements of their prey closely up and down the water column in deep waters. Other species like the megamouth shark adopt a similar feeding strategy that mirrors movement in the water column of their plankton prey. In sub-Arctic lakes, certain morphs of the whitefish (''Coregonus lavaretus'') are planktivorous; the pelagic whitefish feeds primarily on zooplankton and as such have more gill rakers for enhanced feeding than other, non-planktivorous morphs of the same species.


Nutrient limitation in lake systems

The primary limiting nutrient shifts between nitrogen and phosphorus; a resulting consequence of changes in the structure of the food-web, thus limiting primary and secondary production in aquatic ecosystems. The bioavailability of such nutrients drives variation in the biomass and productivity of planktonic species. Due to variance in the N:P excretion of planktivorous fish species, consumer-driven nutrient cycling results in changes in nutrient availability. By feeding on zooplankton, planktivorous fish can increase the rate of nutrient recycling by releasing phosphorus from their prey. Planktivorous fish may release cyanobacteria from nutrient limitation by increasing the concentration of bioavailable phosphorus through excretion. The presence of planktivorous fish can disturb sediments, resulting in an increase in the amount of nutrients that are bioavailable to phytoplankton and further support in phytoplankton nutrient demands.


Planktivore effects on a global scale


Trophic regulation

Planktivory can play an important role in the growth, abundance, and community composition of planktonic species via top-down trophic control. For example, competitive superiority of large zooplankton over smaller species in lake systems leads to large-body dominance in the absence of planktivorous fish as a result of increased food availability and grazing efficiency. Alternatively, the presence of planktivorous fish results in a decrease in zooplankton population through predation and shifts the community composition towards smaller zooplankton by limiting food availability and influencing size-selective predation (see the "
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
" page for more information regarding size-selective predation). Predation by planktivorous fish reduces grazing by zooplankton and subsequently increases phytoplankton primary production and biomass. By limiting the population and growth rate of zooplankton, obligate zooplanktivores are less likely to migrate to the area due to the lack of available food. For example, the presence of gizzard shad in reservoirs has been observed to strongly influence the recruitment of other planktivores. Variations of fish recruitment and mortality rates from nutrient limitation have also been noted in lake ecosystems. Piscivory can have similar top-down effects on planktonic species by influencing the community composition of planktivores. The population of planktivorous fish can also be influenced through predation by piscivorous species such as marine mammals and aquatic birds. For example, planktivorous minnows in Lake Gatun experienced a rapid population decline after the introduction of peacock bass (''Cichla ocellaris''). However, a reduced population of planktivorous fish species result in a population increase of another class of planktivores – zooplankton. In lake ecosystems, some fish have been observed to behave first as zooplanktivores then as piscivores, affecting cascading trophic interactions. Planktivory pressure from zooplankton in marine communities (top-down control, as previously mentioned)has a large influence on phytoplankton productivity. Zooplankton can control phytoplankton seasonal dynamics as they exert the largest grazing pressure on phytoplankton; they also may modify their grazing strategies depending on environmental conditions, leading to seasonal change. For instance, copepods can switch between ambushing prey and using water flow to capture prey depending on external conditions and prey abundance. The planktivorous pressure zooplankton exert could explain the diversity of phytoplankton despite many phytoplankton occupying similar ecological niches (see the " paradox of the plankton" page for more information regarding this ecological conundrum). One notable example of trophic control is how planktivores have the ability to impact the species distribution of larval crabs in estuaries and coastal waters. Crab larvae, which are also planktivores, are hatched inside estuaries but some species then begin their migration out to waters along the coast where there are not as many predators. These crab larvae then utilize the tides to return to the estuaries when they become benthic organisms and are no longer planktivores. Planktivores tend to live their early lives within estuaries. These juvenile fish tend to inhabit these regions throughout the warmer months in the year. Throughout the year, the risk for plankton varies within estuaries, the risk reaches its highest from August to October, and the lowest from December to April, this is consistent with the theory that planktivory is the highest in the summer months in this system. The risk of planktivory is strongly correlated with the number of planktivores within this system.


Nutrient transport

Consumers can regulate primary production in an ecosystem by altering ratios of nutrients via different rates of recycling. Nutrient transport is greatly influenced by planktivorous fish, which recycle and transport nutrients between benthic and pelagic habitats. Nutrients released by benthic-feeding fishes can increase the total nutrient content of pelagic waters, as transported nutrients are fundamentally different from those that are recycled. Additionally, planktivorous fish can have significant effect on nutrient transport as well as total nutrient concentration by disturbing sediments through
bioturbation Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a ...
. Increased nutrient cycling from near-sediment bioturbation by filter-feeding planktivores can increase phytoplankton population via nutrient enrichment. Salmon accumulate marine nutrients as they mature in ocean environments which they then transport back to their stream of origin to spawn. As they decompose, the freshwater streams become enriched with nutrients which contribute to the development of the ecosystem. The physical transport of nutrients and plankton can greatly affect the community composition and food web structure within oceanic ecosystems. In nearshore regions, planktivores and piscivores have been shown to be highly sensitive to changes in ocean currents while zooplankton populations are unable to tainted levels of predation pressure.


Planktivore modification on plankton growth

In some marine systems, planktivory can be an important factor controlling the duration and extent of phytoplankton blooms. Changes in phytoplankton communities and growth rates can modify the amount of grazing pressure present; grazing pressure can also be dampened by physical factors in the water column. The scientist Michael Behrenfeld proposed that the deepening of the
mixed layer The oceanic or limnological mixed layer is a layer in which active turbulence has homogenized some range of depths. The surface mixed layer is a layer where this turbulence is generated by winds, surface heat fluxes, or processes such as evaporat ...
in the ocean, a vertical region near the surface made physically and chemically homogenous by active mixing, leads to a decrease in grazing interactions among planktivores and plankton because planktivores and plankton become more spatially distant from one another. This spatial distance thereby facilitates phytoplankton blooms and ultimately grazing rates by planktivores; both the physical changes and changes to grazing pressure have a significant influence on where and when phytoplankton blooms occur. The shallowing of the mixed layer due to physical processes within the water column conversely intensifies planktivore feeding.


Harmful algal blooms

Harmful algal bloom A harmful algal bloom (HAB), or excessive algae growth, sometimes called a red tide in marine environments, is an algal bloom that causes negative impacts to other organisms by production of natural algae-produced toxins, water deoxygenation, ...
s occur when there is a bloom of toxin producing phytoplankton. Planktivores such as fish and filter feeders that are present have a high likelihood of consuming these phytoplankton because that is what makes up the majority of their diet, or the diet of their prey. Since these planktivores near the bottom of the food chain consume harmful toxins, those toxins then move up the food web when predators consume these fish. The increasing concentration of some toxins through trophic levels presented here is called
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 faster than it can be lost or eliminated by catabolism and excretion. T ...
, and this can lead to a range of impacts from non-lethal changes in behavior to major die-offs of large marine animals. There are monitoring programs in place for shellfish due to human health concerns and the ease of sampling in oysters. Some fish feed directly on phytoplankton, like the
Atlantic herring Atlantic herring (''Clupea harengus'') is a herring in the family Clupeidae. It is one of the most abundant fish species in the world. Atlantic herrings can be found on both sides of the northern Atlantic Ocean, congregating in large schools. ...
(''Clupea harengus''), and
Clupeidae Clupeidae is a family of clupeiform ray-finned fishes, comprising, for instance, the herrings and sprats. Many members of the family have a body protected with shiny cycloid (very smooth and uniform) scales, a single dorsal fin, and a fusiform ...
, while other fish feed on zooplankton that consume the harmful algae.
Domoic acid Domoic acid (DA) is a kainic acid-type neurotoxin that causes amnesic shellfish poisoning (ASP). It is produced by algae and accumulates in shellfish, sardines, and anchovies. When sea lions, otters, cetaceans, humans, and other predators eat cont ...
is a toxin carried by a type of
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
called ''
Pseudo-nitzschia ''Pseudo-nitzschia'' is a marine planktonic diatom genus that accounts for 4.4% of pennate diatoms found worldwide. Some species are capable of producing the neurotoxin domoic acid (DA), which is responsible for the neurological disorder in humans ...
''. Pseudo-nitzchia were the main organism responsible for a large HAB that took place along the west coast of the US in 2015 and had a large impact on the Dungeness crab fishery that year. When harmful algal blooms occur, planktivorous fish can act as vectors for poisonous substances like domoic acid. These planktivorous fish are eaten by larger fish and birds and the subsequent ingestion of toxins can then harm those species. Those animals consume planktivorous fish during a harmful algal bloom, and can have miscarriages, seizures, vomiting, and can sometimes die. Additionally, marine mammal mortality is occasionally attributed to harmful algal blooms, according to NOAA. Krill are another example of a planktivore that may exhibit high levels of domoic acid in their system; these large plankton are then consumed by humpback and blue whales. Since krill can have such a high level of domoic acid in their system when blooms are present, that concentration is rapidly transferred to whales which leads them to have a high concentration of domoic acid in their system as well. There is no evidence proving that this domoic acid has had a negative impact on the whales, but if the concentration of domoic acid is great enough, they could be impacted similarly to other marine mammals.


The role of climate change

Climate change is a worldwide phenomenon that affects everything from the largest planktivores such as whales, to even the smallest plankton. Climate change affects weather patterns, creates seasonal anomalies, alters
sea surface temperature Sea surface temperature (or ocean surface temperature) is the ocean temperature, temperature of ocean water close to the surface. The exact meaning of ''surface'' varies in the literature and in practice. It is usually between and below the sea ...
, alters ocean currents, and can affect nutrient availability for phytoplankton, and may even spur HABs in some systems.


Arctic and Antarctic

The Arctic has been hit hard with shorter winters and hotter summers creating less permafrost and rapidly melting ice caps causing lower salinity levels. The coupling of higher ocean CO2 levels, temperatures, and lower salinity is causing changes in phytoplankton communities and diatom diversity. ''
Thalassiosira ''Thalassiosira'' is a genus of centric diatoms, comprising over 100 marine and freshwater species. It is a diverse group of photosynthetic eukaryotes that make up a vital part of marine and freshwater ecosystems, in which they are key primary pr ...
spp''. Plankton was replaced by solitary ''Cylindrotheca closterium'' or ''
Pseudo-nitzschia ''Pseudo-nitzschia'' is a marine planktonic diatom genus that accounts for 4.4% of pennate diatoms found worldwide. Some species are capable of producing the neurotoxin domoic acid (DA), which is responsible for the neurological disorder in humans ...
spp''., a common HAB causing phytoplankton, under higher temperature and lower salinity in combination. Community changes such as this one, have large-scale effects through trophic levels. A shift in the primary producer communities can cause shifts in consumer communities, as the new food may provide different dietary benefits. As there is less permanent ice in the Arctic and less summer ice, some planktivores species are already moving north into these new open waters.
Atlantic cod The Atlantic cod (: cod; ''Gadus morhua'') is a fish of the family Gadidae, widely consumed by humans. It is also commercially known as '' cod'' or ''codling''.orcas The orca (''Orcinus orca''), or killer whale, is a toothed whale and the largest member of the oceanic dolphin family. The only extant species in the genus '' Orcinus'', it is recognizable by its black-and-white-patterned body. A cosmopoli ...
have been documented in these new territories, while planktivores such as Arctic cod are losing their habitat and feeding grounds under and around the sea ice. Similarly, the Arctic birds, the Least and Crested Auklets rely on zooplankton that lives under the disappearing sea ice and has seen dramatic effects on reproductive fitness and nutrition stress with the decreasing amounts of zooplankton available in the Bering Sea basin. In another prime example of shifting food webs, Moore et al. (2018) have found a shift from benthic dominated ecosystem to a more pelagic dominated ecosystem feeding structure. With longer open water periods, due to a loss of sea ice the
Chukchi Sea The Chukchi Sea (, ), sometimes referred to as the Chuuk Sea, Chukotsk Sea or the Sea of Chukotsk, is a marginal sea of the Arctic Ocean. It is bounded on the west by the Long Strait, off Wrangel Island, and in the east by Point Barrow, Alaska, ...
has seen a shift in the past three decades. The increase in air temperature and loss of sea ice have coupled to promote an increase in pelagic fishes and a decrease in benthic biomass. This shift has encouraged a shift to planktivorous seabirds instead of piscivorous seabirds. Pollock fish are a planktivorous fish that rely on copepods as their primary diet as juveniles. According to the Oscillating Control Hypothesis, early ice retreat caused by a warming climate creates a later bloom of copepods and aphids (a plankton species). The later bloom produces fewer large lipid rich copepods, and results in smaller less nutrient rich copepods. The older pollock then face a winter starvation, causing carnivory on young pollock (<1yr old), and reduced population numbers and fitness. Similar to the Arctic, sea ice in the Antarctic is melting rapidly and permanent ice is becoming less and less (Zachary Lab Cite). This ice melt creates changes in freshwater input and 
ocean stratification Ocean stratification is the natural separation of an ocean's water into horizontal layers by Density of water, density. This is generally stable stratification, because warm water floats on top of cold water, and heating is mostly from the sun, whi ...
, consequently affecting nutrient delivery to primary producers. As sea ice recedes, there is less valuable surface area for algae to grow on the bottom of the ice. This lack of algae inhibits krill (a partial planktonic species) to have less food availability, consequently affecting the fitness of Antarctic primary consumers such as krill, squid, pollock, and other
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...
zooplankton.


Subarctic

The Subarctic has seen similar ecosystem changes especially in well studied places such as Alaska. The warmer waters have helped increase zooplankton communities and have been creating a shift in ecosystem dynamics (Green 2017). There has been a large shift from piscivorous seabirds such as pacific loons and black-legged kittiwakes to planktivores sea birds such as ancient auklets and short-tailed shearwaters. Marine planktivores such as the charismatic humpback,
fin A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. F ...
, and minke whales have been benefiting from the increase in zooplankton such as an increase in krill. As these large whales spend more time migrating into these northern water, they are taking up resources previously only used by arctic planktivores, creating potential shifts in food availability and thus food webs.


Tropics

Tropical and equatorial marine regions are mainly characterized by coral reef communities or vast open oceans.
Coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in group ...
s are one of the most susceptible ecosystems to climate change, in particular the symptoms of warming oceans and acidification.
Ocean acidification Ocean acidification is the ongoing decrease in the pH of the Earth's ocean. Between 1950 and 2020, the average pH of the ocean surface fell from approximately 8.15 to 8.05. Carbon dioxide emissions from human activities are the primary cause of ...
raises CO2 levels in the ocean and has significant effects on zooplankton communities. Smith et al. (2016) discovered that increased levels of CO2 show reductions in zooplankton biomass but not zooplankton quality ''in tropical ecosystems'', as increased CO2 had no negative effects on fatty acid compositions. This means that planktivores are not receiving less nutritious zooplankton, but are experiencing lesser availability of zooplankton than is needed for survival. One of the most important planktivores in the tropics are corals themselves. Although spending a portion of their life cycle as planktonic organisms themselves, established corals are sedentary organisms that can use their tentacles to capture plankton from the surrounding environment to help supplement energy produced by the photosynthetic
zooxanthellae Zooxanthellae (; zooxanthella) is a colloquial term for single-celled photosynthetic organisms that are able to live in symbiosis with diverse marine invertebrates including corals, jellyfish, demosponges, and nudibranchs. Most known zooxanthell ...
. Climate change has had significant impacts on coral reefs, with warming causing coral bleaching and increases in infectious diseases, sea-level rise causing more sedimentation that then smothers corals, stronger and more frequent storms causing breakage and structural destruction, an increase of land runoff bringing more nutrients into the systems causing algal blooms that murk up the water and therefore diminish light availability for photosynthesis, altered ocean currents causing a difference in the dispersal of larvae and planktonic food availability, and lastly changes in ocean pH decreasing structural integrity and growth rates. There is also a plethora of planktivorous fish throughout the tropics that play important ecological roles within marine systems. Similar to corals, planktivorous reef fish are directly affected by these changing systems and these negative effects then disrupt food webs through the oceans. As plankton communities shift in speciation and availability, primary consumers have a harder time meeting energy budgets. This lack of food availability can influence reproductivity and overall primary consumer populations, creating food shortages for higher trophic consumers.


Effects of planktivores on industry

The global
fisheries Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life or, more commonly, the site where such enterprise takes place ( a.k.a., fishing grounds). Commercial fisheries include wild fisheries and fish farm ...
industry is a multi-billion dollar, international industry that provides food and livelihoods to billions of people around the globe. Some of the most important fisheries include salmon, pollock, mackerel, char, cod, halibut, and trout. In 2021, the take home total profits, before bonuses, actually going into fishermen's pockets, from the Alaskan salmon, cod, flounder, and groundfish fishing season came to $248 million. Planktivorous fish alone create an important, large economic industry. In 2017 Alaska pollock was the United States' largest commercial fishery by volume with 3.4 billion pounds being caught and coming in at total value of $413 million. Besides fishing, planktivorous marine animals drive tourism economy as well. Tourist travel across the world for
whale watching Whale watching is the practice of observing whales and dolphins (cetaceans) in their natural habitat. Whale watching is mostly a recreational activity (cf. birdwatching), but it can also serve scientific and/or educational purposes.Hoyt, E. ...
, to see charismatic megafauna such as
humpback whale The humpback whale (''Megaptera novaeangliae'') is a species of baleen whale. It is a rorqual (a member of the family Balaenopteridae) and is the monotypic taxon, only species in the genus ''Megaptera''. Adults range in length from and weigh u ...
s in Hawaii, Minke whales in Alaska, grey whales in Oregon, and
whale shark The whale shark (''Rhincodon typus'') is a slow-moving, filter feeder, filter-feeding carpet shark and the largest known Extant taxon, extant fish species. The largest confirmed individual had a length of . The whale shark holds many records for ...
s in South America.
Manta ray Manta rays are large Batoidea, rays belonging to the genus ''Mobula'' (formerly its own genus ''Manta''). The larger species, ''Giant oceanic manta ray, M. birostris'', reaches in width, while the smaller, ''Reef manta ray, M. alfredi'', reac ...
s also drive dive and snorkel tourism, raking in over $73 million annually, in direct revenue, over 23 countries around the world. The main participating countries in Manta ray tourism include Japan, Indonesia, the Maldives, Mozambique, Thailand, Australia, Mexico, United States, Federated States of Micronesia and Palau.


See also

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Zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
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Plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
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Piscivore A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
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Phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
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Filter feeder Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a s ...
*
Carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...


References

{{feeding Ecology terminology Animals by eating behaviors Limnology