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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 water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
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 w ...
s. Zooplankton can be contrasted with phytoplankton, which are the plant component of the plankton community ("phyto" comes from the Greek word for ''plant''). Zooplankton are
heterotrophic 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 ...
(other-feeding), whereas phytoplankton are
autotrophic An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
(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 or microscope to see them clearly. In physics, the microscopic scale is sometimes regarded as the scale be ...
, 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 umbrell ...
) 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. Histo ...
(single-celled protists that prey on other microscopic life) are zooplankton, including zooflagellates,
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 for catching food and other uses; and commonly ...
ns,
radiolarian The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ectoplasm. The elab ...
s, some dinoflagellates and marine microanimals. Macroscopic zooplankton include pelagic cnidarians, ctenophores, molluscs,
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
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 ...
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
mixotroph 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 ...
s. 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)
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
. The word ''zooplankton'' is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''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 protozoans 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 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 their lif ...
ic organisms whose complete
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring * Life-cycle hypothesis ...
lies within the plankton, as well as meroplanktonic organisms that spend part of their lives in the plankton before graduating to either the nekton or a Sessility (zoology), sessile, benthos, benthic existence. Although zooplankton are primarily transported by ambient water currents, many have Animal locomotion, locomotion, used to avoid predators (as in diel vertical migration) 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 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 levels (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 (element), mercury. File:Typical ocean models featuring zooplankton 2.jpg,

Ecology, 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 for catching food and other uses; and commonly ...
ns, radiolarians and dinoflagellates (the last of these are often
mixotroph 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 ...
ic). Important metazoan zooplankton include cnidarians 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 umbrell ...
and the Portuguese Man o' War; crustaceans such as cladocerans, copepods, ostracods, isopods, amphipods, mysids and krill; Chaetognatha, chaetognaths (arrow worms); Mollusca, molluscs such as pteropods; and chordates such as salps and juvenile fish. This wide phylogeny, phylogenetic range includes a similarly wide range in feeding behavior: filter feeder, filter feeding, predation and symbiosis with
autotrophic An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
phytoplankton as seen in corals. Zooplankton feed on bacterioplankton, phytoplankton, other zooplankton (sometimes Cannibalism (zoology), cannibalistically), detritus (or marine snow) and even Nekton, 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 infectious disease, disease natural reservoir, reservoir. Crustacean zooplankton have been found to house the bacterium ''Vibrio cholerae'', which causes cholera, by allowing the cholera vibrios to attach to their chitinous exoskeletons. This symbiotic 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 Morphology (biology), 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, 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 level, 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 ectotherms 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 Zooplankton grazing, plankton...'' Microzooplankton are defined as heterotrophic and Mixotrophic dinoflagellate, mixotrophic plankton. They primarily consist of phagotrophic protists, including ciliates, dinoflagellates, and mesozooplankton Nauplius (larva), nauplii. 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 creativecommons:by/4.0/, 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

Protozoans are protists that feed on organic matter such as other microorganisms or organic tissues and debris. Historically, the protozoa were regarded as "one-celled animals", because they often possess animal-like behaviours, such as motility and predation, and lack a cell wall, as found in plants and many algae. 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 zooflagellates,
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 for catching food and other uses; and commonly ...
ns,
radiolarian The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ectoplasm. The elab ...
s and some dinoflagellates.


Radiolarians

Radiolarians are unicellular predatory #Marine protists, protists 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 #Marine microfossils, 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 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 for catching food and other uses; and commonly ...
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 Test (biology), tests, are chambered (forams add more chambers as they grow). The shells are usually made of calcite, but are sometimes made of Agglutination (biology), agglutinated sediment particles or chiton, 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'' 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 pyramids 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'' sp. File:Collection Penard MHNG Specimen 533-2-1 Pamphagus granulatus.tif, wiktionary:xenogenic, Xenogenic testate amoeba covered in diatoms


Ciliates

File:Stylonychia putrina - 160x - II (13215594964).jpg, ''Stylonychia, Stylonychia putrina'' File:Holophyra ovum - 400x (9836710085).jpg, ''Holophyra'' ovum File:Mikrofoto.de-Blepharisma japonicum 15.jpg, ''Blepharisma japonicum'' File:The ciliate Frontonia sp.jpg, This ciliate is digesting cyanobacteria. The mouth is at the bottom right.


Dinoflagellates

Dinoflagellates are a phylum of unicellular flagellates with about 2,000 marine species. Some dinoflagellates are Predatory dinoflagellate, predatory, 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. Excavata, Excavates may be the most basal flagellate lineage. File:Gyrodinium dinoflagellate.jpg, ''Gymnodinium, 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 radiolarians can be in symbiosis with dinoflagellates Dinoflagellates often live in symbiosis with other organisms. Many nassellarian radiolarians house dinoflagellate Symbiosis, symbionts within their tests. The nassellarian provides ammonium and carbon dioxide 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 for catching food and other uses; and commonly ...
. File:Ceratium tripos.jpg, Tripos (dinoflagellate), ''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'' produces red tides highly toxic to humans File:Algal bloom(akasio) by Noctiluca in Nagasaki.jpg, Red tide


Mixotrophs

A
mixotroph 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 ...
is an organism that can use a mix of different Primary nutritional groups, 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 chloroplasts, 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 phototroph, photo- and chemotrophy, lithotroph, litho- and organotrophy, autotroph, 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 eukaryote, eukaryotic or prokaryote, 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 (ecology), biomass was mixotrophic. ''Phaeocystis'' species are endosymbionts to Acantharea, acantharian radiolarians. ''Phaeocystis'' is an important algal genus found as part of the marine phytoplankton around the world. It has a Polymorphism (biology), 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 Algal bloom, blooms. As a result, ''Phaeocystis'' is an important contributor to the marine Carbon cycle, carbon and sulfur cycles. File:Tintinnid ciliate Favella.jpg, Tintinnid ciliate ''Favella'' File:Euglena mutabilis - 400x - 1 (10388739803) (cropped).jpg, ''Euglena, Euglena mutabilis'', a photosynthetic flagellate File:Stichotricha secunda - 400x (14974779356).jpg, Zoochlorellae (green) living inside the ciliate ''Stichotricha secunda'' File:Dinophysis acuta.jpg, The dinoflagellate ''Dinophysis acuta'' A number of forams are mixotrophic. These have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates. Mixotrophic foraminifers are particularly common in nutrient-poor oceanic waters. Some forams are kleptoplasty, kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis. By trophic orientation dinoflagellates are all over the place. Some dinoflagellates are known to be photosynthesis, photosynthetic, but a large fraction of these are in fact mixotrophy, mixotrophic, 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 reefs. Others predate other protozoa, and a few forms are parasitic. Many dinoflagellates are mixotrophic and could also be classified as phytoplankton. The toxic dinoflagellate ''Dinophysis acuta'' acquire chloroplasts from its prey. "It cannot catch the cryptophytes by itself, and instead relies on ingesting ciliates such as the red ''Myrionecta rubra'', which sequester their chloroplasts from a specific cryptophyte clade (Geminigera/Plagioselmis/Teleaulax)".


Metazoa (animals)

Copepods 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 made of calcium carbonate and usually have a Crustacean larvae#Nauplius, 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. File:Copepod 2 with eggs.jpg, Copepod with eggs File:Tomopteriskils.jpg, Segmented worm File:Hyperia.jpg, Amphipod File:Krill666.jpg, Krill File:Glaucus atlanticus 1 cropped.jpg, Glaucus atlanticus, Blue ocean slug


Holoplankton and meroplankton


Ichthyoplankton

Ichthyoplankton are the Fish eggs, eggs and larvae 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 File:Molalavdj.jpg, Ocean sunfish larvae (2.7mm) File:FMIB 47039 Ostracion hoops.jpeg, Boxfish larva


Gelatinous zooplankton

Gelatinous zooplankton include ctenophores, Jellyfish, medusae, salps, and Chaetognatha in coastal waters. Jellyfish are slow swimmers, and most species form part of the plankton. Traditionally jellyfish have been viewed as Trophic level, trophic 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 and the leatherback sea turtle.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.
Graeme Hays, 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 salps and ctenophores, 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, Tetrapturus, spearfish and swordfish as well as various birds and invertebrates such as octopus, sea cucumbers, crabs and amphipods. "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 siphonophores, ctenophores and cephalopods. File:Parumbrosa polylobata 01.jpg, Jellyfish File:Ctenophora-Comb-jelly-Euplokamis-sp.-02.jpg, Bioluminescent ctenophore (comb jelly) File:Tunicate off Atauro island.jpg, This free-floating pyrosome is made up of hundreds of individual bioluminescent tunicates File:23 salpchain frierson odfw (8253212250).jpg, Salp 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.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 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. 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 cycle, 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 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 Mesopelagic zone, twilight zone.


Role in biogeochemistry

In addition to linking primary producers to higher trophic levels in marine food webs, zooplankton also play an important role as “recyclers” of carbon and other nutrients that significantly impact marine biogeochemical cycles, including the biological pump. This is particularly important in the oligotrophic waters of the open ocean. Through sloppy feeding, excretion, egestion, and leaching of fecal pellets, zooplankton release dissolved organic matter (DOM) which controls DOM cycling and supports the microbial loop. 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 Marine carbon export, carbon export, including the production of fecal pellets, mucous feeding webs, molts, and carcasses. Zooplankton fecal pellets, Fecal pellets 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 falls – 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 Benthos, benthic organisms.


See also

* Census of Marine Zooplankton * Diel vertical migration * Ocean acidification * Primary production * Thin layers (oceanography)


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