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Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal'').
Plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by currents in the
ocean The ocean (also the sea or the world ocean) is the body of salt water that covers approximately 70.8% of the surface of Earth and contains 97% of Earth's water. An ocean can also refer to any of the large bodies of water into which the wo ...
, or by currents in
sea The sea, connected as the world ocean or simply the ocean, is the body of salty water that covers approximately 71% of the Earth's surface. The word sea is also used to denote second-order sections of the sea, such as the Mediterranean Sea, ...
s,
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s or
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of wate ...
s. Zooplankton can be contrasted with
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'. Ph ...
, which are the plant component of the plankton community ("phyto" comes from the Greek word for ''plant''). Zooplankton are heterotrophic (other-feeding), whereas phytoplankton are autotrophic (self-feeding). This means zooplankton cannot manufacture their own food but must eat other plants or animals instead — in particular they eat phytoplankton. Zooplankton are generally larger than phytoplankton, most are
microscopic The microscopic scale () is the scale of objects and events smaller than those that can easily be seen by the naked eye, requiring a lens (optics), lens or microscope to see them clearly. In physics, the microscopic scale is sometimes regarded a ...
, but some (such as
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
) are macroscopic and can be seen with the
naked eye Naked eye, also called bare eye or unaided eye, is the practice of engaging in visual perception unaided by a magnifying, light-collecting optical instrument, such as a telescope or microscope, or eye protection. Vision corrected to normal ...
. Many
protozoans Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histor ...
(single-celled
protists A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the excl ...
that prey on other microscopic life) are zooplankton, including
zooflagellate In some older systems of classification, Zoomastigophora is a phylum (more commonly known as zooflagellates) within the kingdom Protista. Organisms within this group have a spherical, elongated body with a single central nucleus. They are single-ce ...
s,
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
ns, radiolarians, some
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s and
marine microanimals Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...
. Macroscopic zooplankton include pelagic
cnidarian Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
s,
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
s,
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
s,
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s and
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 ca ...
s, as well as planktonic
arrow worms The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and can ...
and
bristle worms Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made ...
. The distinction between plants and animals often breaks down in very small organisms. Recent studies of marine microplankton have indicated over half of microscopic plankton are mixotrophs. A mixotroph is an organism that can behave sometimes as though it were a plant, and sometimes as though it were an animal, using a mix of autotrophy and heterotrophy. Many marine microzooplankton are mixotrophic, which means they could also be classified as phytoplankton.


Overview

Zooplankton (; ) are
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic (sometimes
detritivorous Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrat ...
)
plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
. The word ''zooplankton'' is derived from the Greek ''zoon'' (), meaning "animal", and ' (), meaning "wanderer" or "drifter". Zooplankton is a categorization spanning a range of
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
sizes including small
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
ns and large
metazoa Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
ns. It includes
holoplankton Holoplankton are organisms that are plankton, planktic (they live in the water column and cannot swim against a current) for their entire life cycle. Holoplankton can be contrasted with meroplankton, which are planktic organisms that spend part of ...
ic organisms whose complete life cycle lies within the plankton, as well as
meroplankton Meroplankton are a wide variety of aquatic organisms which have both planktonic and benthic stages in their life cycles. Much of the meroplankton consists of larval stages of larger organism. Meroplankton can be contrasted with holoplankton, whic ...
ic organisms that spend part of their lives in the plankton before graduating to either the nekton or a
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
,
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
existence. Although zooplankton are primarily transported by ambient water currents, many have locomotion, used to avoid predators (as in
diel vertical migration Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The word ''diel'' comes from the Latin ''dies'' day, and means a 24-h ...
) or to increase prey encounter rate. Just as any species can be limited within a geographical region, so are zooplankton. However, species of zooplankton are not dispersed uniformly or randomly within a region of the ocean. As with phytoplankton, ‘patches’ of zooplankton species exist throughout the ocean. Though few physical barriers exist above the mesopelagic, specific species of zooplankton are strictly restricted by salinity and temperature gradients; while other species can withstand wide temperature and salinity gradients. Zooplankton patchiness can also be influenced by biological factors, as well as other physical factors. Biological factors include breeding, predation, concentration of phytoplankton, and vertical migration. The physical factor that influences zooplankton distribution the most is mixing of the water column (
upwelling Upwelling is an oceanographic phenomenon that involves wind-driven motion of dense, cooler, and usually nutrient-rich water from deep water towards the ocean surface. It replaces the warmer and usually nutrient-depleted surface water. The nutr ...
and downwelling along the coast and in the open ocean) that affects nutrient availability and, in turn, phytoplankton production. Through their consumption and processing of phytoplankton and other food sources, zooplankton play a role in aquatic food webs, as a resource for consumers on higher
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
s (including fish), and as a conduit for packaging the organic material in the biological pump. Since they are typically small, zooplankton can respond rapidly to increases in phytoplankton abundance, for instance, during the spring bloom. Zooplankton are also a key link in the biomagnification of pollutants such as
mercury Mercury commonly refers to: * Mercury (planet), the nearest planet to the Sun * Mercury (element), a metallic chemical element with the symbol Hg * Mercury (mythology), a Roman god Mercury or The Mercury may also refer to: Companies * Merc ...
. File:Typical ocean models featuring zooplankton 2.jpg,

Ecologically important protozoan zooplankton groups include the
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
ns, radiolarians and
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s (the last of these are often mixotrophic). Important metazoan zooplankton include
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
ns such as
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
and the
Portuguese Man o' War The Portuguese man o' war (''Physalia physalis''), also known as the man-of-war, is a marine hydrozoan found in the Atlantic Ocean and the Indian Ocean. It is considered to be the same species as the Pacific man o' war or blue bottle, which is ...
;
crustacea Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gro ...
ns such as cladocerans,
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 (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s,
ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typic ...
s,
isopod Isopoda is an order of crustaceans that includes woodlice and their relatives. Isopods live in the sea, in fresh water, or on land. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, an ...
s,
amphipod Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far descr ...
s,
mysids Mysida is an order of small, shrimp-like crustaceans in the malacostracan superorder Peracarida. Their common name opossum shrimps stems from the presence of a brood pouch or "marsupium" in females. The fact that the larvae are reared in this ...
and krill; chaetognaths (arrow worms); molluscs such as pteropods; and
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 fiv ...
s such as
salp A salp (plural salps, also known colloquially as “sea grape”) or salpa (plural salpae or salpas) is a barrel-shaped, planktic tunicate. It moves by contracting, thereby pumping water through its gelatinous body, one of the most efficient ...
s and juvenile fish. This wide
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
range includes a similarly wide range in feeding behavior: filter feeding,
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
and
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
with autotrophic
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'. Ph ...
as seen in corals. Zooplankton feed on
bacterioplankton Bacterioplankton refers to the bacterial component of the plankton that drifts in the water column. The name comes from the Ancient Greek word ' ('), meaning "wanderer" or "drifter", and ', a Latin term coined in the 19th century by Christian G ...
, phytoplankton, other zooplankton (sometimes cannibalistically),
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
(or marine snow) and even nektonic organisms. As a result, zooplankton are primarily found in surface waters where food resources (phytoplankton or other zooplankton) are abundant. Zooplankton can also act as a
disease A disease is a particular abnormal condition that negatively affects the structure or function of all or part of an organism, and that is not immediately due to any external injury. Diseases are often known to be medical conditions that a ...
reservoir A reservoir (; from French ''réservoir'' ) is an enlarged lake behind a dam. Such a dam may be either artificial, built to store fresh water or it may be a natural formation. Reservoirs can be created in a number of ways, including contro ...
. Crustacean zooplankton have been found to house the bacterium ''
Vibrio cholerae ''Vibrio cholerae'' is a species of Gram-negative, facultative anaerobe and comma-shaped bacteria. The bacteria naturally live in brackish or saltwater where they attach themselves easily to the chitin-containing shells of crabs, shrimps, and oth ...
'', which causes
cholera Cholera is an infection of the small intestine by some strains of the bacterium ''Vibrio cholerae''. Symptoms may range from none, to mild, to severe. The classic symptom is large amounts of watery diarrhea that lasts a few days. Vomiting and ...
, by allowing the cholera vibrios to attach to their chitinous
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
s. This
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
relationship enhances the bacterium's ability to survive in an aquatic environment, as the exoskeleton provides the bacterium with carbon and nitrogen.


Size classification

Body size has been defined as a "master trait" for plankton as it is a morphological characteristic shared by organisms across taxonomy that characterises the functions performed by organisms in ecosystems. It has a paramount effect on growth, reproduction, feeding strategies and mortality. One of the oldest manifestations of the biogeography of traits was proposed over 170 years ago, namely
Bergmann's rule Bergmann's rule is an ecogeographical rule that states that within a broadly distributed taxonomic clade, populations and species of larger size are found in colder environments, while populations and species of smaller size are found in warmer ...
, in which field observations showed that larger species tend to be found at higher, colder latitudes. Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
In the oceans, size is critical in determining trophic links in planktonic ecosystems and is thus a critical factor in regulating the efficiency of the biological carbon pump. Body size is sensitive to changes in temperature due to the thermal dependence of physiological processes. The plankton is mainly composed of
ectotherm An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...
s which are organisms that do not generate sufficient metabolic heat to elevate their body temperature, so their metabolic processes depends on external temperature. Consequently, ectotherms grow more slowly and reach maturity at a larger body size in colder environments, which has long puzzled biologists because classic theories of life-history evolution predict smaller adult sizes in environments delaying growth. This pattern of body size variation, known as the temperature-size rule (TSR), has been observed for a wide range of ectotherms, including single-celled and multicellular species, invertebrates and vertebrates. The processes underlying the inverse relationship between body size and temperature remain to be identified. Despite temperature playing a major role in shaping latitudinal variations in organism size, these patterns may also rely on complex interactions between physical, chemical and biological factors. For instance, oxygen supply plays a central role in determining the magnitude of ectothermic temperature-size responses, but it is hard to disentangle the relative effects of oxygen and temperature from field data because these two variables are often strongly inter-related in the surface ocean. Zooplankton can be broken down into size classes which are diverse in their morphology, diet, feeding strategies, etc. both within classes and between classes:


Picozooplankton

<2μm


Nanozooplankton

2-20μm


Microzooplankton

20-200μm ''Major grazers of the
plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
...'' Microzooplankton are defined as heterotrophic and
mixotrophic A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comp ...
plankton. They primarily consist of phagotrophic
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s, including ciliates, dinoflagellates, and
mesozooplankton 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 ...
nauplii Nauplius, Nauplia or Nauplios , may refer to : Greece-related * Nauplius (mythology), in Greek mythology, the son of Poseidon and Amymone, the father of Palamedes, and also the name of an Argonaut * Nauplia Nafplio ( ell, Ναύπλιο) is a ...
. As the primary consumers of marine phytoplankton, microzooplankton consume ~ 59–75% daily of the marine primary production, much larger than mesozooplankton. That said, macrozooplankton can sometimes have greater consumption rates in eutrophic ecosystems because the larger phytoplankton can be dominant there. Microzooplankton are also pivotal regenerators of nutrients which fuel primary production and food sources for metazoans. Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License. Despite their ecological importance, microzooplankton remain understudied. Routine oceanographic observations seldom monitor microzooplankton biomass or herbivory rate, although the dilution technique, an elegant method of measuring microzooplankton herbivory rate, has been developed for almost four decades (Landry and Hassett 1982). The number of observations of microzooplankton herbivory rate is around 1600 globally, far less than that of primary productivity (> 50,000). This makes validating and optimizing the grazing function of microzooplankton difficult in ocean ecosystem models.


Mesozooplankton

0.2-20 millimeters Because plankton is rarely fished, it has been argued that mesoplankton abundance and species composition can be used to study marine ecosystems' response to climate change. This is because they have life cycles that generally last less than a year, meaning they respond to climate changes between years. Sparse, monthly sampling will still indicate vacillations.


Taxonomic groups


Protozoans

Protozoan Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
s are
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s that feed on organic matter such as other
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s or organic tissues and debris. Historically, the protozoa were regarded as "one-celled animals", because they often possess
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 ...
-like behaviours, such as
motility Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
and
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
, and lack a
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
, as found in plants and many
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
. Although the traditional practice of grouping protozoa with animals is no longer considered valid, the term continues to be used in a loose way to identify single-celled organisms that can move independently and feed by
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
y. Marine protozoans include
zooflagellate In some older systems of classification, Zoomastigophora is a phylum (more commonly known as zooflagellates) within the kingdom Protista. Organisms within this group have a spherical, elongated body with a single central nucleus. They are single-ce ...
s,
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
ns, radiolarians and some
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s.


Radiolarians

Radiolarians are unicellular predatory
protists A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the excl ...
encased in elaborate globular shells usually made of silica and pierced with holes. Their name comes from the Latin for "radius". They catch prey by extending parts of their body through the holes. As with the silica frustules of diatoms, radiolarian shells can sink to the ocean floor when radiolarians die and become preserved as part of the ocean sediment. These remains, as microfossils, provide valuable information about past oceanic conditions.Wassilieff, Maggy (2006
"Plankton - Animal plankton"
''Te Ara - the Encyclopedia of New Zealand''. Accessed: 2 November 2019.
File:Mikrofoto.de-Radiolarien 6.jpg, Like diatoms, radiolarians come in many shapes File:Podocyrtis papalis Ehrenberg - Radiolarian (30448963206).jpg, Also like diatoms, radiolarian shells are usually made of silicate File:Acantharian radiolarian Xiphacantha (Haeckel).jpg, However acantharian radiolarians have shells made from
strontium sulfate Strontium sulfate (SrSO4) is the sulfuric acid, sulfate salt of strontium. It is a white crystalline powder and occurs in nature as the mineral Celestine (mineral), celestine. It is poorly soluble in water to the extent of 1 part in 8,800. It is mo ...
crystals File:Spherical radiolarian 2.jpg, Cutaway schematic diagram of a spherical radiolarian shell


Foraminiferans

Like radiolarians,
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
ns (''forams'' for short) are single-celled predatory protists, also protected with shells that have holes in them. Their name comes from the Latin for "hole bearers". Their shells, often called
tests Test(s), testing, or TEST may refer to: * Test (assessment), an educational assessment intended to measure the respondents' knowledge or other abilities Arts and entertainment * ''Test'' (2013 film), an American film * ''Test'' (2014 film), ...
, are chambered (forams add more chambers as they grow). The shells are usually made of calcite, but are sometimes made of agglutinated sediment particles or
chiton Chitons () are marine molluscs of varying size in the class Polyplacophora (), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized. They are also sometimes known as gumboots or sea cradles or coat-of-mail s ...
, and (rarely) of silica. Most forams are benthic, but about 40 species are planktic. They are widely researched with well established fossil records which allow scientists to infer a lot about past environments and climates. File:EB1911 Foraminifera - Section of Rotalia beccarii.jpg, section showing chambers of a spiral foram File:Live Ammonia tepida.jpg, Live ''
Ammonia tepida ''Ammonia tepida'' is a benthic foraminifer living in the sediment of brackish waters. It is very similar to ''Ammonia beccarii'', but the latter lives on the surface of red algae. Once considered a globally widespread taxon, a recent genetic an ...
'' streaming granular ectoplasm for catching food File:Planktic Foraminifera of the northern Gulf of Mexico.jpg, Group of planktonic forams File:All Gizah Pyramids.jpg, The
Egyptian pyramid The Egyptian pyramids are ancient masonry structures located in Egypt. Sources cite at least 118 identified "Egyptian" pyramids. Approximately 80 pyramids were built within the Kingdom of Kush, now located in the modern country of Sudan. Of ...
s were constructed from limestone that contained nummulites.


Amoeba

File:Amoeba proteus 2.jpg, Naked amoeba sketch showing food vacuoles and ingested diatom File:Arcella sp.jpg, Shell or test of a testate amoeba, ''
Arcella ''Arcella'' is a genus of testate amoebae in the order Arcellinida, usually found in freshwaters and mosses, and rarely in soils. A key characteristic of ''Arcella'' is the circular test with a hole on its center from where finger-like pseudopod ...
'' sp. File:Collection Penard MHNG Specimen 533-2-1 Pamphagus granulatus.tif,
Xenogenic A xenobiotic is a chemical substance found within an organism that is not naturally produced or expected to be present within the organism. It can also cover substances that are present in much higher concentrations than are usual. Natural compo ...
testate amoeba covered in diatoms


Ciliates

File:Stylonychia putrina - 160x - II (13215594964).jpg, '' Stylonychia putrina'' File:Holophyra ovum - 400x (9836710085).jpg, ''Holophyra'' ovum File:Mikrofoto.de-Blepharisma japonicum 15.jpg, ''
Blepharisma japonicum ''Blepharisma japonicum'' is a species of protozoan that can be found either in water or soil in Japan. Description The body of an organism is elongated and ovoided. It has a curved anterior apex that is over the peristome. It has a cytostome ...
'' File:The ciliate Frontonia sp.jpg, This
ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
is digesting
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
. The mouth is at the bottom right.


Dinoflagellates

Dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s are a phylum of unicellular
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
s with about 2,000 marine species. Some dinoflagellates are
predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
, and thus belong to the zooplankton community. Their name comes from the Greek "dinos" meaning ''whirling'' and the Latin "flagellum" meaning a ''whip'' or ''lash''. This refers to the two whip-like attachments (flagella) used for forward movement. Most dinoflagellates are protected with red-brown, cellulose armour.
Excavates Excavation may refer to: * Excavation (archaeology) * Excavation (medicine) * ''Excavation'' (The Haxan Cloak album), 2013 * ''Excavation'' (Ben Monder album), 2000 * ''Excavation'' (novel), a 2000 novel by James Rollins * '' Excavation: A Memo ...
may be the most basal flagellate lineage. File:Gyrodinium dinoflagellate.jpg, '' Gyrodinium'', one of the few naked dinoflagellates which lack armour File:Protoperidinium dinoflagellate.jpg, The dinoflagellate ''Protoperidinium'' extrudes a large feeding veil to capture prey File:Radiolarian - Podocyrtis (Lampterium) mitra Ehrenberg - 160x.jpg,
Nassellarian Nassellaria is an order of Rhizaria belonging to the class Radiolaria. The organisms of this order are characterized by a skeleton cross link with a cone or ring. Introduction Nassellaria is an order of Radiolaria under the class Polycystina. ...
radiolarians can be in symbiosis with dinoflagellates
Dinoflagellates often live in
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
with other organisms. Many
nassellarian Nassellaria is an order of Rhizaria belonging to the class Radiolaria. The organisms of this order are characterized by a skeleton cross link with a cone or ring. Introduction Nassellaria is an order of Radiolaria under the class Polycystina. ...
radiolarians house
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
symbionts Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
within their tests. The nassellarian provides
ammonium The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternary a ...
and
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
for the dinoflagellate, while the dinoflagellate provides the nassellarian with a mucous membrane useful for hunting and protection against harmful invaders. There is evidence from DNA analysis that dinoflagellate symbiosis with radiolarians evolved independently from other dinoflagellate symbioses, such as with
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
. File:Ceratium tripos.jpg, ''Tripos muelleri'' is recognisable by its U-shaped horns File:Archives de zoologie expérimentale et générale (1920) (20299351186).jpg, '' Oodinium'', a genus of parasitic dinoflagellates, causes velvet disease in fish File:Karenia brevis.jpg, ''
Karenia brevis ''Karenia brevis'' is a microscopic, single-celled, photosynthetic organism in the genus '' Karenia''. It is a marine dinoflagellate commonly found in the waters of the Gulf of Mexico. It is the organism responsible for the "Florida red tides" t ...
'' produces red tides highly toxic to humans File:Algal bloom(akasio) by Noctiluca in Nagasaki.jpg, Red tide


Mixotrophs

A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comprise more than half of all microscopic plankton. There are two types of eukaryotic mixotrophs: those with their own
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s, and those with endosymbionts—and others that acquire them through kleptoplasty or by enslaving the entire phototrophic cell. The distinction between plants and animals often breaks down in very small organisms. Possible combinations are photo- and chemotrophy, litho- and
organotroph An organotroph is an organism that obtains hydrogen or electrons from organic substrates. This term is used in microbiology to classify and describe organisms based on how they obtain electrons for their respiration processes. Some organotrophs su ...
y, auto- and
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
y or other combinations of these. Mixotrophs can be either
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
or prokaryotic. They can take advantage of different environmental conditions. Many marine microzooplankton are mixotrophic, which means they could also be classified as phytoplankton. Recent studies of marine microzooplankton found 30–45% of the ciliate abundance was mixotrophic, and up to 65% of the amoeboid, foram and radiolarian
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bi ...
was mixotrophic. ''Phaeocystis'' species are endosymbionts to acantharian radiolarians. '' Phaeocystis'' is an important algal genus found as part of the marine
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'. Ph ...
around the world. It has a polymorphic life cycle, ranging from free-living cells to large colonies. It has the ability to form floating colonies, where hundreds of cells are embedded in a gel matrix, which can increase massively in size during blooms. As a result, ''Phaeocystis'' is an important contributor to the marine
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent In chemistry, the valence (US spelling) or valency (British spelling) of an element is the measure of its combining capacity with o ...
and
sulfur cycle The sulfur cycle is a biogeochemical cycle in which the sulfur moves between rocks, waterways and living systems. It is important in geology as it affects many minerals and in life because sulfur is an essential element ( CHNOPS), being a const ...
s. File:Tintinnid ciliate Favella.jpg,
Tintinnid Tintinnids are ciliates of the choreotrich order Tintinnida, distinguished by vase-shaped shells, the name deriving from a Latin source meaning a small tinkling bell, that are called'' loricae'', which are mostly protein but may incorporate min ...
ciliate ''Favella'' File:Euglena mutabilis - 400x - 1 (10388739803) (cropped).jpg, '' Euglena mutabilis'', a photosynthetic
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
File:Stichotricha secunda - 400x (14974779356).jpg,
Zoochlorellae ''Zoochlorella'' is a ''nomen rejiciendum'' for a genus of green algae assigned to ''Chlorella''. The term zoochlorella (plural zoochlorellae) is sometimes used to refer to any green algae that lives symbiotically within the body of a freshwater ...
(green) living inside the
ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
''Stichotricha secunda'' File:Dinophysis acuta.jpg, The dinoflagellate ''Dinophysis acuta''
A number of forams are mixotrophic. These have unicellular
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
as endosymbionts, from diverse lineages such as the
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
,
red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
, golden algae,
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s, and
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s. Mixotrophic foraminifers are particularly common in nutrient-poor oceanic waters. Some forams are kleptoplastic, retaining
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s from ingested algae to conduct
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
. By trophic orientation dinoflagellates are all over the place. Some dinoflagellates are known to be
photosynthetic Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in c ...
, but a large fraction of these are in fact
mixotrophic A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comp ...
, combining photosynthesis with ingestion of prey ( phagotrophy). Some species are endosymbionts of marine animals and other protists, and play an important part in the biology of
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 groups. Co ...
s. Others predate other protozoa, and a few forms are parasitic. Many dinoflagellates are
mixotrophic A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comp ...
and could also be classified as phytoplankton. The toxic dinoflagellate ''
Dinophysis acuta ''Dinophysis acuta'' is a species of flagellated planktons belonging to the genus ''Dinophysis''. It is one of the few unusual photosynthetic protists that acquire plastids from algae by endosymbiosis. By forming massive blooms, particularly ...
'' acquire chloroplasts from its prey. "It cannot catch the cryptophytes by itself, and instead relies on ingesting ciliates such as the red ''
Myrionecta rubra ''Mesodinium rubrum'' (or ''Myrionecta rubra'') is a species of ciliates. It constitutes a plankton community and is found throughout the year, most abundantly in spring and fall, in coastal areas. Although discovered in 1908, its scientific im ...
'', which sequester their chloroplasts from a specific cryptophyte clade (Geminigera/Plagioselmis/Teleaulax)".


Metazoa (animals)

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 (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s are typically 1 to 2 mm long with a teardrop-shaped bodies. Like all crustaceans, their bodies are divided into three sections: head, thorax, and abdomen, with two pairs of antennae; the first pair is often long and prominent. They have a tough
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
made of calcium carbonate and usually have a single red eye in the centre of their transparent head. About 13,000 species of copepods are known, of which about 10,200 are marine. They are usually among the more dominant members of the zooplankton. Metazoan zooplankton" style="float:left"> File:Copepod 2 with eggs.jpg,
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 (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
with eggs File:Tomopteriskils.jpg,
Segmented worm The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
File:Hyperia.jpg,
Amphipod Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far descr ...
File:Krill666.jpg, Krill File:Glaucus atlanticus 1 cropped.jpg, Blue ocean slug


Holoplankton and meroplankton


Ichthyoplankton

Ichthyoplankton Ichthyoplankton (from Greek: ἰχθύς, , "fish"; and πλαγκτός, , "drifter") are the eggs and larvae of fish. They are mostly found in the sunlit zone of the water column, less than 200 metres deep, which is sometimes called the epip ...
are the eggs and
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
of fish ("ichthyo" comes from the Greek word for ''fish''). They are planktonic because they cannot swim effectively under their own power, but must drift with the ocean currents. Fish eggs cannot swim at all, and are unambiguously planktonic. Early stage larvae swim poorly, but later stage larvae swim better and cease to be planktonic as they grow into juvenile fish. Fish larvae are part of the zooplankton that eat smaller plankton, while fish eggs carry their own food supply. Both eggs and larvae are themselves eaten by larger animals. File:Squidu.jpg, Juvenile planktonic
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
File:Molalavdj.jpg,
Ocean sunfish The ocean sunfish or common mola (''Mola mola'') is one of the largest bony fish in the world. It was misidentified as the heaviest bony fish, which was actually a different species, ''Mola alexandrini''. Adults typically weigh between . The spe ...
larvae (2.7mm) File:FMIB 47039 Ostracion hoops.jpeg,
Boxfish Ostraciidae is a family of squared, bony fish belonging to the order Tetraodontiformes, closely related to the pufferfishes and filefishes. Fish in the family are known variously as boxfishes, cofferfishes, cowfishes and trunkfishes. It contains ...
larva


Gelatinous zooplankton

Gelatinous zooplankton Gelatinous zooplankton are fragile animals that live in the water column in the ocean. Their delicate bodies have no hard parts and are easily damaged or destroyed. Gelatinous zooplankton are often transparent. All jellyfish are gelatinous zoopla ...
include
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
s, medusae,
salps A salp (plural salps, also known colloquially as “sea grape”) or salpa (plural salpae or salpas) is a barrel-shaped, planktic tunicate. It moves by contracting, thereby pumping water through its gelatinous body, one of the most efficient ...
, and
Chaetognatha The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and can ...
in coastal waters. Jellyfish are slow swimmers, and most species form part of the plankton. Traditionally jellyfish have been viewed as
trophic Trophic, from Ancient Greek τροφικός (''trophikos'') "pertaining to food or nourishment", may refer to: * Trophic cascade * Trophic coherence * Trophic egg * Trophic function * Trophic hormone * Trophic level index * Trophic level * Trop ...
dead ends, minor players in the marine food web, gelatinous organisms with a body plan largely based on water that offers little nutritional value or interest for other organisms apart from a few specialised predators such as the
ocean sunfish The ocean sunfish or common mola (''Mola mola'') is one of the largest bony fish in the world. It was misidentified as the heaviest bony fish, which was actually a different species, ''Mola alexandrini''. Adults typically weigh between . The spe ...
and the
leatherback sea turtle The leatherback sea turtle (''Dermochelys coriacea''), sometimes called the lute turtle or leathery turtle or simply the luth, is the largest of all living turtles and the heaviest non-crocodilian reptile, reaching lengths of up to and weights ...
.Hamilton, G. (2016
"The secret lives of jellyfish: long regarded as minor players in ocean ecology, jellyfish are actually important parts of the marine food web"
''Nature'', 531(7595): 432-435.
Hays, G.C., Doyle, T.K. and Houghton, J.D. (2018) "A paradigm shift in the trophic importance of jellyfish?" ''Trends in ecology & evolution'', 33(11): 874-884. That view has recently been challenged. Jellyfish, and more gelatinous zooplankton in general, which include
salp A salp (plural salps, also known colloquially as “sea grape”) or salpa (plural salpae or salpas) is a barrel-shaped, planktic tunicate. It moves by contracting, thereby pumping water through its gelatinous body, one of the most efficient ...
s and
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
s, are very diverse, fragile with no hard parts, difficult to see and monitor, subject to rapid population swings and often live inconveniently far from shore or deep in the ocean. It is difficult for scientists to detect and analyse jellyfish in the guts of predators, since they turn to mush when eaten and are rapidly digested. But jellyfish bloom in vast numbers, and it has been shown they form major components in the diets of
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...
,
spearfish Spearfish may refer to: Places *Spearfish, South Dakota, United States * North Spearfish, South Dakota, United States * Spearfish Formation, a geologic formation in the United States Biology * ''Tetrapturus'', a genus of marlin containing spe ...
and
swordfish Swordfish (''Xiphias gladius''), also known as broadbills in some countries, are large, highly migratory predatory fish characterized by a long, flat, pointed bill. They are a popular sport fish of the billfish category, though elusive. Swordfis ...
as well as various birds and invertebrates such as
octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttle ...
,
sea cucumber Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothuria ...
s,
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...
s and
amphipod Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far descr ...
s. "Despite their low energy density, the contribution of jellyfish to the energy budgets of predators may be much greater than assumed because of rapid digestion, low capture costs, availability, and selective feeding on the more energy-rich components. Feeding on jellyfish may make marine predators susceptible to ingestion of plastics." According to a 2017 study, narcomedusae consume the greatest diversity of mesopelagic prey, followed by
physonect Physonectae is a suborder of siphonophores. In Japanese it is called (). Organisms in the suborder Physonectae follow the classic Siphonophore body plan. They are almost all pelagic, and are composed of a colony of specialized zooids that ori ...
siphonophores,
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
s and
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s. File:Parumbrosa polylobata 01.jpg,
Jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
File:Ctenophora-Comb-jelly-Euplokamis-sp.-02.jpg,
Bioluminescent Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorganisms including ...
ctenophore Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
(comb jelly) File:Tunicate off Atauro island.jpg, This free-floating pyrosome is made up of hundreds of individual
bioluminescent Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorganisms including ...
tunicates File:23 salpchain frierson odfw (8253212250).jpg,
Salp A salp (plural salps, also known colloquially as “sea grape”) or salpa (plural salpae or salpas) is a barrel-shaped, planktic tunicate. It moves by contracting, thereby pumping water through its gelatinous body, one of the most efficient ...
chain
The importance of the so-called "jelly web" is only beginning to be understood, but it seems medusae, ctenophores and siphonophores can be key predators in deep pelagic food webs with ecological impacts similar to predator fish and squid. Traditionally gelatinous predators were thought ineffectual providers of marine trophic pathways, but they appear to have substantial and integral roles in deep
pelagic food webs Compared to terrestrial environments, marine environments have biomass pyramids which are inverted at the base. In particular, the biomass of consumers (copepods, krill, shrimp, forage fish) is larger than the biomass of primary producers. This ...
.Choy, C.A., Haddock, S.H. and Robison, B.H. (2017) "Deep pelagic food web structure as revealed by ''in situ'' feeding observations". ''Proceedings of the Royal Society B: Biological Sciences'', 284(1868): 20172116. . Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Role in food webs

Grazing by single-celled zooplankton accounts for the majority of
organic carbon Total organic carbon (TOC) is the amount of carbon found in an organic compound and is often used as a non-specific indicator of water quality or cleanliness of pharmaceutical manufacturing equipment. TOC may also refer to the amount of organic c ...
loss from marine primary production. Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
However, zooplankton grazing remains one of the key unknowns in global predictive models of carbon flux, the marine food web structure and ecosystem characteristics, because empirical grazing measurements are sparse, resulting in poor parameterisation of grazing functions. To overcome this critical knowledge gap, it has been suggested that a focused effort be placed on the development of instrumentation that can link changes in phytoplankton biomass or optical properties with grazing. Grazing is a central, rate-setting process in ocean ecosystems and a driver of
marine biogeochemical cycling Marine biogeochemical cycles are biogeochemical cycles that occur within marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. These biogeochemical cycles are the pathways chemical substan ...
. In all ocean ecosystems, grazing by heterotrophic protists constitutes the single largest loss factor of marine primary production and alters particle size distributions. Grazing affects all pathways of export production, rendering grazing important both for surface and deep carbon processes. Predicting central paradigms of ocean ecosystem function, including responses to environmental change requires accurate representation of grazing in global biogeochemical, ecosystem and cross-biome-comparison models. Several large-scale analyses have concluded that phytoplankton losses, which are dominated by grazing are the putative explanation for annual cycles in phytoplankton biomass, accumulation rates and export production. File:Export Processes in the Ocean from Remote Sensing.jpg,
Pelagic food web Compared to terrestrial environments, marine environments have biomass pyramids which are inverted at the base. In particular, the biomass of consumers (copepods, krill, shrimp, forage fish) is larger than the biomass of primary producers. Thi ...
and the biological pump. Links among the ocean's biological pump and pelagic food web and the ability to sample these components remotely from ships, satellites, and autonomous vehicles. Light blue waters are the euphotic zone, while the darker blue waters represent the twilight zone.


Role in biogeochemistry

In addition to linking primary producers to higher
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
s in marine food webs, zooplankton also play an important role as “recyclers” of carbon and other nutrients that significantly impact
marine biogeochemical cycles Marine biogeochemical cycles are biogeochemical cycles that occur within marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. These biogeochemical cycles are the pathways chemical substan ...
, including the biological pump. This is particularly important in the
oligotrophic An oligotroph is an organism that can live in an environment that offers very low levels of nutrients. They may be contrasted with copiotrophs, which prefer nutritionally rich environments. Oligotrophs are characterized by slow growth, low rates of ...
waters of the open ocean. Through sloppy feeding, excretion, egestion, and leaching of fecal pellets, zooplankton release
dissolved organic matter Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particu ...
(DOM) which controls DOM cycling and supports the
microbial loop The microbial loop describes a trophic pathway where, in aquatic systems, dissolved organic carbon (DOC) is returned to higher trophic levels via its incorporation into bacterial biomass, and then coupled with the classic food chain formed by phy ...
. Absorption efficiency, respiration, and prey size all further complicate how zooplankton are able to transform and deliver carbon to the deep ocean.


Sloppy feeding and release of DOM

Excretion and sloppy feeding (the physical breakdown of food source) make up 80% and 20% of crustacean zooplankton-mediated DOM release respectively. In the same study, fecal pellet leaching was found to be an insignificant contributor. For protozoan grazers, DOM is released primarily through excretion and egestion and gelatinous zooplankton can also release DOM through the production of mucus. Leaching of fecal pellets can extend from hours to days after initial egestion and its effects can vary depending on food concentration and quality. Various factors can affect how much DOM is released from zooplankton individuals or populations. Absorption efficiency (AE) is the proportion of food absorbed by plankton that determines how available the consumed organic materials are in meeting the required physiological demands. Depending on the feeding rate and prey composition, variations in AE may lead to variations in fecal pellet production, and thus regulates how much organic material is recycled back to the marine environment. Low feeding rates typically lead to high AE and small, dense pellets, while high feeding rates typically lead to low AE and larger pellets with more organic content. Another contributing factor to DOM release is respiration rate. Physical factors such as oxygen availability, pH, and light conditions may affect overall oxygen consumption and how much carbon is loss from zooplankton in the form of respired CO2. The relative sizes of zooplankton and prey also mediate how much carbon is released via sloppy feeding. Smaller prey are ingested whole, whereas larger prey may be fed on more “sloppily”, that is more biomatter is released through inefficient consumption. There is also evidence that diet composition can impact nutrient release, with carnivorous diets releasing more dissolved organic carbon (DOC) and ammonium than omnivorous diets.


Carbon export

Zooplankton play a critical role in supporting the ocean's biological pump through various forms of carbon export, including the production of fecal pellets, mucous feeding webs, molts, and carcasses.
Fecal pellets Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
are estimated to be a large contributor to this export, with copepod size rather than abundance expected to determine how much carbon actually reaches the ocean floor. The importance of fecal pellets can vary both by time and location. For example, zooplankton bloom events can produce larger quantities of fecal pellets, resulting in greater measures of carbon export. Additionally, as fecal pellets sink, they are reworked by microbes in the water column, which can thus alter the carbon composition of the pellet. This affects how much carbon is recycled in the euphotic zone and how much reaches depth. Fecal pellet contribution to carbon export is likely underestimated; however, new advances in quantifying this production are currently being developed, including the use of isotopic signatures of amino acids to characterize how much carbon is being exported via zooplankton fecal pellet production. Carcasses are also gaining recognition as being important contributors to carbon export.
Jelly fall Jelly-falls are marine carbon cycling events whereby gelatinous zooplankton, primarily cnidarians, sink to the seafloor and enhance carbon and nitrogen fluxes via rapidly sinking particulate organic matter. These events provide nutrition to benth ...
s – the mass sinking of gelatinous zooplankton carcasses – occur across the world as a result of large blooms. Because of their large size, these gelatinous zooplankton are expected to hold a larger carbon content, making their sinking carcasses a potentially important source of food for
benthic organisms Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.Census of Marine Zooplankton The Census of Marine Zooplankton is a field project of the Census of Marine Life that has aimed to produce a global assessment of the species diversity, biomass, biogeographic distribution, and genetic diversity of more than 7,000 described specie ...
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Diel vertical migration Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The word ''diel'' comes from the Latin ''dies'' day, and means a 24-h ...
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Ocean acidification Ocean acidification is the reduction in the pH value of the Earth’s ocean. Between 1751 and 2021, the average pH value of the ocean surface has decreased from approximately 8.25 to 8.14. The root cause of ocean acidification is carbon dioxid ...
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Primary production In ecology, primary production is the synthesis of organic compounds from atmospheric or aqueous carbon dioxide. It principally occurs through the process of photosynthesis, which uses light as its source of energy, but it also occurs through c ...
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Thin layers (oceanography) Thin layers are concentrated aggregations of phytoplankton and zooplankton in coastal and offshore waters that are vertically compressed to thicknesses ranging from several centimeters up to a few meters and are horizontally extensive, sometime ...


References


External links


SAHFOS
Sir Alister Hardy Foundation for Ocean Science
Ocean Drifters
Short film narrated by David Attenborough about the varied roles of plankton
Sea Drifters
BBC Audio slideshow
Plankton Chronicles
Short documentary films & photos
COPEPOD: The global plankton database
A global coverage database of zooplankton biomass and abundance data.
Guide to the marine zooplankton of south eastern AustraliaTasmanian Aquaculture and Fisheries Institute




{{Authority control Aquatic animals Biological oceanography Planktology Oceanographical terminology de:Plankton#Zooplankton