Krill

Krill are small crustaceans of the order Euphausiacea, and are found in all the
Euphausiacea
world's oceans. The name "krill" comes from the Norwegian word krill, meaning
"small fry of fish",[1] which is also often attributed to species of fish.

Krill are considered an important trophic level connection – near the bottom of
the food chain – because they feed on phytoplankton and (to a lesser extent)
zooplankton, converting these into a form suitable for many larger animals for
which krill make up the largest part of their diets. In the Southern Ocean, one
species, the Antarctic krill, Euphausia superba, makes up an estimatedbiomass of
Northern krill (Meganyctiphanes
around 379,000,000 tonnes,[2] making it among the species with the largest total
biomass. Of this, over half is eaten by whales, seals, penguins, squid, and fish norvegica)
each year, and is replaced by growth and reproduction. Most krill species display Scientific classification
large daily vertical migrations, thus providing food for predators near the surface
Kingdom: Animalia
at night and in deeper waters during the day.
Phylum: Euarthropoda
Krill are fished commercially in the Southern Ocean and in the waters around
Japan. The total global harvest amounts to 150,000–200,000 tonnes annually,
Subphylum: Crustacea
most of this from the Scotia Sea. Most of the krill catch is used for aquaculture Class: Malacostraca
and aquarium feeds, as bait in sport fishing, or in the pharmaceutical industry. In
Superorder: Eucarida
Japan, the Philippines, and Russia, krill are also used for human consumption and
are known as okiami (オキアミ) in Japan. They are eaten as camarones in Spain Order: Euphausiacea
and Philippines. In the Philippines, krill are also known as alamang and are used Dana, 1852
to make a salty paste calledbagoong.
Families and genera
Krill are also the main prey ofbaleen whales, including the blue whale.
Euphausiidae

Euphausia Dana, 1852

Contents
Meganyctiphanes Holt and W.
Taxonomy
Phylogeny M. Tattersall , 1905
Timeline Nematobrachion Calman , 1905
Distribution
Nematoscelis G. O. Sars , 1883
Anatomy and morphology
Nyctiphanes G. O. Sars, 1883
Ecology
Feeding Pseudeuphausia Hansen , 1910
Predation
Plastics Stylocheiron G. O. Sars, 1883
Life history and behavior Tessarabrachion Hansen, 1911
Reproduction
Thysanoessa Brandt, 1851
Moulting
Lifespan Thysanopoda Latreille , 1831
Swarming
Vertical migration Bentheuphausiidae
Human uses
Harvesting history
Human consumption
 See also Bentheuphausia G. O. Sars,
References
1885
Further reading
External links

Taxonomy
Krill belong to the large arthropod subphylum, the Crustacea. The most familiar and largest group of crustaceans, the class
Malacostraca, includes the superorder Eucarida comprising the three orders, Euphausiacea (krill), Decapoda (shrimp, prawns,
lobsters, crabs), and the planktonicAmphionidacea.

The order Euphausiacea comprises two families. The more abundant Euphausiidae contains 10 different genera with a total of 85
species. Of these, the genus Euphausia is the largest, with 31 species.[3] The lesser known family, the Bentheuphausiidae, has only
one species, Bentheuphausia amblyops, a bathypelagic krill living in deep waters below 1,000 m (3,300 ft). It is considered the most
primitive extant krill species.[4]

Well-known species of the Euphausiidae of commercial krill fisheries include Antarctic krill (Euphausia superba), Pacific krill (E.
pacifica) and Northern krill (Meganyctiphanes norvegica).[5]

Phylogeny
Proposed phylogeny of Euphausiacea[6]
Bentheuphausiidae Bentheuphausia

Thysanopoda (♣)

Nematobrachion (♦)

Meganyctiphanes

Pseudeuphausia
Euphausiini (♠)(♦)
Euphausia
Euphausiacea
Euphausiidae
Nyctiphanes
Euphausiinae
Nematoscelis

Nematoscelini (♠) Thysanoessa

Nematoscelina (♠)
Tessarabrachion

Stylocheiron

Phylogeny obtained from morphological data, (♠) names coined in,[6] (♣) possibly paraphyletic taxon
due to Nematobrachion in.[6] (♦) clades differs from Casanova (1984),[7] where Pseudoeuphausia is
sister to Nyctiphanes, Euphausia is sister to Thysanopoda and Nematobrachion is sister to
Stylocheiron.
As of 2013, the order Euphausiacea is believed to be monophyletic due to several unique conserved morphological characteristics
[8] and by molecular studies.[9][10][11]
(autapomorphy) such as its naked filamentous gills and thin thoracopods
There have been many theories of the location of the order Euphausiacea. Since the first description of Thysanopode tricuspide by
Henri Milne-Edwards in 1830, the similarity of their biramous thoracopods had led zoologists to group euphausiids and Mysidacea in
the order Schizopoda, which was split by Johan Erik Vesti Boas in 1883 into two separate orders.[12] Later, William Thomas Calman
(1904) ranked the Mysidacea in the superorder Peracarida and euphausiids in the superorder Eucarida, although even up to the 1930s
the order Schizopoda was advocated.[8] It was later also proposed that order Euphausiacea should be grouped with the Penaeidae
(family of prawns) in the Decapoda based on developmental similarities, as noted by Robert Gurney and Isabella Gordon.[13][14] The
[8]
reason for this debate is that krill share some morphological features of decapods and others of mysids.

Molecular studies have not unambiguously grouped them, possibly due to the paucity of key rare species such as Bentheuphausia
amblyops in krill and Amphionides reynaudii in Eucarida. One study supports the monophyly of Eucarida (with basal Mysida),[15]
[10] while yet another groups Euphausiacea withHoplocarida.[16]
another groups Euphausiacea with Mysida (the Schizopoda),

Timeline
No extant fossil can be unequivocally assigned to Euphausiacea. Some extinct eumalacostracan taxa have been thought to be
euphausiaceans such asAnthracophausia, Crangopsis – now assigned to the Aeschronectida (Hoplocarida)[6] – and Palaeomysis.[17]
All dating of speciation events were estimated by molecular clock methods, which placed the last common ancestor of the krill
family Euphausiidae (order Euphausiacea minus Bentheuphausia amblyops) to have lived in the Lower Cretaceous about
130 million years ago.[10]

Distribution
Krill occur worldwide in all oceans, although many individual species have endemic or neritic (i.e., coastal) distributions.
Bentheuphausia amblyops, a bathypelagic species, has a cosmopolitan distributionwithin its deep-sea habitat.[18]

Species of the genus Thysanoessa occur in both Atlantic and Pacific oceans.[19] The Pacific is home to Euphausia pacifica. Northern
krill occur across the Atlantic from theMediterranean Sea northward.

Species with neritic distributions include the four species of the genusNyctiphanes.[20] They are highly abundant along the upwelling
regions of the California, Humboldt, Benguela, and Canarias current systems.[21][22][23] Another species having only neritic
distribution is E. crystallorophias, which is endemic to the Antarctic coastline.[24]

Species with endemic distributions include Nyctiphanes capensis, which occurs only in the Benguela current,[20] E. mucronata in the
Humboldt current,[25] and the six Euphausia species native to the Southern Ocean.

In the Antarctic, seven species are known,[26] one in genus Thysanoessa (T. macrura) and six in Euphausia. The Antarctic krill
(Euphausia superba) commonly lives at depths reaching 100 m (330 ft),[27] whereas ice krill (Euphausia crystallorophias) reach
depth of 4,000 m (13,100 ft), though they commonly inhabit depths of at most 300–600 m (1,000–2,000 ft).[28] Both are found at
latitudes south of 55° S, with E. crystallorophias dominating south of 74° S[29] and in regions of pack ice. Other species known in the
Southern Ocean are E. frigida, E. longirostris, E. triacantha and E. vallentini.[30]

Anatomy and morphology

Krill are crustaceans and have a chitinous exoskeleton made up of three tagmata: the cephalon (head), the pereion (fused to the
cephalon to form a cephalothorax), and the pleon. This outer shell of krill is transparent in most species. Krill feature intricate
compound eyes; some species adapt to different lighting conditions through the use of screening pigments.[31] They have two
antennae and several pairs of thoracic legs called pereiopods or thoracopods, so named because they are attached to the thorax; their
number varies among genera and species. These thoracic legs include feeding legs and grooming legs. Additionally all species have
five pairs of swimming legs called pleopods or "swimmerets", very similar to those of a lobster or freshwater crayfish. Most krill are
about 1–2 centimetres (0.4–0.8 in) long as adults; a few species grow to sizes on the order of 6–15 centimetres (2.4–5.9 in). The
largest krill species is the bathypelagic Thysanopoda spinicauda.[32] Krill can be easily distinguished from other crustaceans such as
true shrimp by their externally visiblegills.[33]
Except for Bentheuphausia amblyops, krill are bioluminescent animals having
organs called photophores that can emit light. The light is generated by an enzyme-
catalysed chemiluminescence reaction, wherein a luciferin (a kind of pigment) is
activated by a luciferase enzyme. Studies indicate that the luciferin of many krill
species is a fluorescent tetrapyrrole similar but not identical to dinoflagellate
luciferin[34] and that the krill probably do not produce this substance themselves but
acquire it as part of their diet, which contains dinoflagellates.[35] Krill photophores
are complex organs with lenses and focusing abilities, and can be rotated by
Krill anatomy explained, using
muscles.[36] The precise function of these organs is as yet unknown; possibilities Euphausia superba as a model
include mating, social interaction or orientation and as a form of counter-
illumination camouflage to compensate their shadow against overhead ambient
light.[37][38]

Ecology

Feeding
Many krill are filter feeders:[22] their frontmost appendages, the thoracopods, form
very fine combs with which they can filter out their food from the water. These
The gills of krill are externally visible
filters can be very fine indeed in those species (such as Euphausia spp.) that feed
primarily on phytoplankton, in particular on diatoms, which are unicellular algae.
Krill are mostly omnivorous,[39] although a few species arecarnivorous, preying on small zooplankton and fish larvae.[40]

Krill are an important element of the aquatic food chain. Krill convert the primary production of their prey into a form suitable for
consumption by larger animals that cannot feed directly on the minuscule algae. Northern krill and some other species have a
relatively small filtering basket and actively huntcopepods and larger zooplankton.[40]

Predation
Many animals feed on krill, ranging from smaller animals likefish or penguins to larger ones like seals and baleen whales.[41]

Disturbances of an ecosystem resulting in a decline in the krill population can have far-reaching effects. During a coccolithophore
bloom in the Bering Sea in 1998,[42] for instance, the diatom concentration dropped in the affected area. Krill cannot feed on the
smaller coccolithophores, and consequently the krill population (mainly E. pacifica) in that region declined sharply. This in turn
affected other species: the shearwater population dropped. The incident was thought to have been one reason salmon did not spawn
that season.[43]

Climate change poses another threat to krill populations.[44] Several single-celled endoparasitoidic ciliates of the genus Collinia can
infect species of krill and devastate affected populations. Such diseases were reported for Thysanoessa inermis in the Bering Sea and
also for E. pacifica, Thysanoessa spinifera, and T. gregaria off the North American Pacific coast.[45][46] Some ectoparasites of the
family Dajidae (epicaridean isopods) afflict krill (and also shrimp and mysids); one such parasite is Oculophryxus bicaulis, which
was found on the krillStylocheiron affine and S. longicorne. It attaches itself to the animal's eyestalkand sucks blood from its head; it
flicted animals reached maturity.[47]
apparently inhibits the host's reproduction, as none of the af

Plastics
Preliminary research indicates krill can digest microplastics under 5 mm (0.20 in) in diameter, breaking them down and excreting
them back into the environment in smaller form.[48]

Life history and behavior

The life cycle of krill is relatively well understood, despite minor variations in detail
from species to species.[13][22] After krill hatch, they experience several larval
stages—nauplius, pseudometanauplius, metanauplius, calyptopsis, and furcilia, each
of which divides into sub-stages. The pseudometanauplius stage is exclusive to
species that lay their eggs within an ovigerous sac: so-called "sac-spawners". The
larvae grow and moult repeatedly as they develop, replacing their rigid exoskeleton
when it becomes too small. Smaller animals moult more frequently than larger ones.
Yolk reserves within their body nourish the larvae through metanauplius stage. By
the calyptopsis stages differentiation has progressed far enough for them to develop
A nauplius of Euphausia pacifica
a mouth and a digestive tract, and they begin to eat phytoplankton. By that time their
hatching, emerging backwards from
yolk reserves are exhausted and the larvae must have reached the photic zone, the the egg
upper layers of the ocean where algae flourish. During the furcilia stages, segments
with pairs of swimmerets are added, beginning at the frontmost segments. Each new
pair becomes functional only at the next moult. The number of segments added during any one of the furcilia stages may vary even
within one species depending on environmental conditions.[49] After the final furcilia stage, an immature juvenile emerges in a shape
similar to an adult, and subsequently developsgonads and matures sexually.[50]

Reproduction
During the mating season, which varies by species and climate, the male deposits a
sperm sack at the female's genital opening (named thelycum). The females can carry
several thousand eggs in their ovary, which may then account for as much as one
third of the animal's body mass.[51] Krill can have multiple broods in one season,
[23][52]
with interbrood intervals lasting on the order of days.

The head of a female krill of the sac-
spawning species Nematoscelis
difficilis with her brood sac. The eggs
have a diameter of 0.3–0.4
millimetres (0.012–0.016 in)
Krill employ two types of spawning mechanism.[23] The 57 species of the genera
Bentheuphausia, Euphausia, Meganyctiphanes, Thysanoessa, and Thysanopoda are
"broadcast spawners": the female releases the fertilised eggs into the water, where
they usually sink, disperse, and are on their own. These species generally hatch in
the nauplius 1 stage, but have recently been discovered to hatch sometimes as
metanauplius or even as calyptopis stages.[53] The remaining 29 species of the other
genera are "sac spawners", where the female carries the eggs with her, attached to
the rearmost pairs of thoracopods until they hatch as metanauplii, although some
species like Nematoscelis difficilismay hatch as nauplius or pseudometanauplius.[54]

Moulting
Moulting occurs whenever a specimen outgrows its rigid exoskeleton. Young animals, growing faster, moult more often than older
and larger ones. The frequency of moulting varies widely by species and is, even within one species, subject to many external factors
such as latitude, water temperature, and food availability. The subtropical species Nyctiphanes simplex, for instance, has an overall
inter-moult period of two to seven days: larvae moult on the average every four days, while juveniles and adults do so, on average,
every six days. For E. superba in the Antarctic sea, inter-moult periods ranging between 9 and 28 days depending on the temperature
between −1 and 4 °C (30 and 39 °F) have been observed, and for Meganyctiphanes norvegica in the North Sea the inter-moult
periods range also from 9 and 28 days but at temperatures between 2.5 and 15 °C (36.5 and 59.0 °F).[55] E. superba is able to reduce
its body size when there is not enough food available, moulting also when its exoskeleton becomes too large.[56] Similar shrinkage
has also been observed for E. pacifica, a species occurring in the Pacific Ocean from polar to temperate zones, as an adaptation to
[57]
abnormally high water temperatures. Shrinkage has been postulated for other temperate-zone species of krill as well.

Lifespan
Some high-latitude species of krill can live for more than six years (e.g., Euphausia superba); others, such as the mid-latitude species
Euphausia pacifica, live for only two years.[5] Subtropical or tropical species' longevity is still shorter, e.g., Nyctiphanes simplex,
which usually lives for only six to eight months.[58]

Swarming
Most krill are swarming animals; the sizes and densities of such swarms vary by
species and region. For Euphausia superba, swarms reach 10,000 to 60,000
individuals per cubic meter.[59][60] Swarming is a defensive mechanism, confusing
smaller predators that would like to pick out individuals. In 2012, Gandomi and
Alavi presented what appears to be a successful stochastic algorithm for modelling
the behaviour of krill swarms. The algorithm is based on three main factors: " (i)
movement induced by the presence of other individuals (ii) foraging activity, and
(iii) random diffusion."[61]
A krill swarm

Vertical migration
Krill typically follow a diurnal vertical migration. It has been assumed that they
spend the day at greater depths and rise during the night toward the surface. The
deeper they go, the more they reduce their activity,[62] apparently to reduce
encounters with predators and to conserve energy. Swimming activity in krill varies
with stomach fullness. Sated animals that had been feeding at the surface swim less
actively and therefore sink below the mixed layer.[63] As they sink they produce
feces which implies a role in the Antarctic carbon cycle. Krill with empty stomachs
swim more actively and thus head towards the surface. Vertical migration may be a
2–3 times daily occurrence. Some species (e.g., Euphausia superba, E. pacifica, E.
Beating pleopods of a swimming
hanseni, Pseudeuphausia latifrons, and Thysanoessa spinifera) form surface swarms
Antarctic krill
during the day for feeding and reproductive purposes even though such behaviour is
[64]
dangerous because it makes them extremely vulnerable to predators.

Experimental studies using Artemia salina as a model suggest that the vertical migrations of krill several hundreds of metres, in
ef on ocean mixing.[65]
groups tens of metres deep, could collectively create enough downward jets of water to have a significantfect

Dense swarms can elicit a feeding frenzy among fish, birds and mammal predators, especially near the surface. When disturbed, a
, leaving the exuvia behind as a decoy.[66]
swarm scatters, and some individuals have even been observed to moult instantaneously

Krill normally swim at a pace of 5–10 cm/s (2–3 body lengths per second),[67] using their swimmerets for propulsion. Their larger
migrations are subject to ocean currents. When in danger, they show an escape reaction called lobstering – flicking their caudal
structures, the telson and the uropods, they move backwards through the water relatively quickly, achieving speeds in the range of 10
to 27 body lengths per second, which for large krill such as E. superba means around 0.8 m/s (3 ft/s).[68] Their swimming
performance has led many researchers to classify adult krill as micro-nektonic life-forms, i.e., small animals capable of individual
[69]
motion against (weak) currents. Larval forms of krill are generally considered zooplankton.

Human uses

Harvesting history
Krill have been harvested as a food source for humans and domesticated animals since at least the 19th century, and possibly earlier
in Japan, where it was known as okiami. Large-scale fishing developed in the late 1960s and early 1970s, and now occurs only in
Antarctic waters and in the seas around Japan. Historically, the largest krill fishery nations were Japan and the Soviet Union, or, after
the latter's dissolution, Russia and Ukraine.[70] The harvest peaked, which in 1983
was about 528,000 tonnes in the Southern Ocean alone (of which the Soviet Union
[71]
took in 93%), is now managed as a precaution against overfishing.

In 1993, two events caused a decline in krill fishing: Russia exited the industry; and
the Convention for the Conservation of Antarctic Marine Living Resources
(CCAMLR) defined maximum catch quotas for a sustainable exploitation of
Antarctic krill. After an October 2011 review, the Commission decided not to
change the quota.[72] Deep frozen plates of Antarctic krill
for use as animal feed and raw
The annual Antarctic catch stabilised at around 100,000 tonnes, which is roughly material for cooking
one fiftieth of the CCAMLR catch quota.[73] The main limiting factor was probably
high costs along with political and legal issues.[74] The Japanese fishery saturated at
some 70,000 tonnes.[75]

Although krill are found worldwide, fishing in Southern Oceans are preferred because the krill are more "catchable" and abundant in
these regions. Particularly in Antarctic seas which are considered aspristine, they are considered a "clean product".[70]

In 2018 it was announced that almost every krill fishing company operating in Antarctica will abandon operations in huge areas
fer zones" around breeding colonies of penguins.[76]
around the Antarctic Peninsula from 2020, including "buf

Human consumption
Although the total biomass of Antarctic krill may be as abundant as 400 million
tonnes, the human impact on this keystone species is growing, with a 39% increase
in total fishing yield to 294,000 tonnes over 2010–2014.[73] Major countries
involved in krill harvesting areNorway (56% of total catch in 2014), the Republic of
Korea (19%), and China (18%).[73]

Krill is a rich source of protein and omega-3 fatty acids which are under
development in the early 21st Century as human food, dietary supplements as oil
capsules, livestock food, and pet food.[70][72][77] Krill tastes salty with a somewhat
Bagoong alamang, a type of shrimp
stronger fish flavor than shrimp. For mass-consumption and commercially prepared
paste from the Philippines made with
products, they must be peeled to remove the inedibleexoskeleton.[77]
fermented krill

In 2011, the US Food and Drug Administrationpublished a letter of no objection for

a manufactured krill oil product to be generally recognized as safe (GRAS) for
human consumption.[78]

Krill (and other planktonic shrimp) are most widely consumed in Southeast Asia, where it is fermented (with the shells intact) and
usually ground finely to make shrimp paste. It can be stir-fried and eaten paired with white rice or used to add umami flavors to a
wide variety of traditional dishes.[79][80] The liquid from the fermentation process is also harvested asfish sauce.[81]

See also
Crustaceans portal
Cold-water shrimp

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Further reading
Boden, Brian P.; Johnson, Martin W.; Brinton, Edward: "Euphausiacea (Crustacea) of the North Pacific"
. Bulletin of
the Scripps Institution of Oceanography. Volume 6 Number 8, 1955.
Brinton, Edward: "Euphausiids of Southeast Asian waters". Naga Report volume 4, part 5. La Jolla: University of
California, Scripps Institution of Oceanography
, 1975.
Conway, D. V. P.; White, R. G.; Hugues-Dit-Ciles, J.; Galienne, C. .P; Robins, D. B.: Guide to the coastal and surface
zooplankton of the South-Western Indian Ocean, Order Euphausiacea, Occasional Publication of theMarine
Biological Association of the United KingdomNo. 15, Plymouth, UK, 2003.
Everson, I. (ed.): Krill: biology, ecology and fisheries. Oxford, Blackwell Science; 2000.ISBN 0-632-05565-0.
Hamner, William M. (May 1984). "Krill — Untapped Bounty From the Sea?".National Geographic. Vol. 165 no. 5.
pp. 626–642. ISSN 0027-9358. OCLC 643483454.
Mauchline, J.: Euphausiacea: Adults, Conseil International pour l'Exploration de la Mer
, 1971. Identification sheets
for adult krill with many line drawings.PDF file, 2 Mb.
Mauchline, J.: Euphausiacea: Larvae, Conseil International pour l'Exploration de la Mer
, 1971. Identification sheets
for larval stages of krill with many line drawings. PDF file, 3 Mb.
Tett, P.: The biology of Euphausiids, lecture notes from a 2003 course in Marine Biologyfrom Napier University.
Tett, P.: Bioluminescence, lecture notes from the 1999/2000 edition of that same course.

External links
Webcam of Krill Aquarium at Australian Antarctic Division
'Antarctic Energies' animation by Lisa Roberts
Retrieved from "https://en.wikipedia.org/w/index.php?title=Krill&oldid=885471819
"

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