Krill

Euphausiacea
A northern krill (Meganyctiphanes norvegica)
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Crustacea
Class:	Malacostraca
Superorder:	Eucarida
Order:	Euphausiacea Dana, 1852
Families and genera
Euphausiidae Euphausia Dana, 1852 Meganyctiphanes Holt and W. M. Tattersall, 1905 Nematobrachion Calman, 1905 Nematoscelis G. O. Sars, 1883 Nyctiphanes G. O. Sars, 1883 Pseudeuphausia Hansen, 1910 Stylocheiron G. O. Sars, 1883 Tessarabrachion Hansen, 1911 Thysanoessa Brandt, 1851 Thysanopoda Latreille, 1831 Bentheuphausiidae Bentheuphausia amblyops G. O. Sars, 1883

Krill is the common name given to the order Euphausiacea of shrimp-like marine crustaceans. Also known as euphausiids, these small invertebrates are found in all oceans of the world. The common name krill comes from the Norwegian word krill, meaning "young fry of fish",^[1] which is also often attributed to other 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 whom krill makes up the largest part of their diet. In the Southern Ocean, one species, the Antarctic krill, Euphausia superba, makes up an estimated biomass of over 500,000,000 tonnes (490,000,000 long tons; 550,000,000 short tons), roughly twice that of humans. Of this, over half is eaten by whales, seals, penguins, squid and fish each year, and is replaced by growth and reproduction. Most krill species display large daily vertical migrations, thus providing food for predators near the surface at night and in deeper waters during the day.

Commercial fishing of krill is done in the Southern Ocean and in the waters around Japan. The total global harvest amounts to 150,000–200,000 tonnes (150,000–200,000 long tons; 170,000–220,000 short tons) annually, most of this from the Scotia Sea. Most of the krill catch is used for aquaculture and aquarium feeds, as bait in sport fishing, or in the pharmaceutical industry. In Japan and Russia, krill is also used for human consumption and is known as okiami (オキアミ^?) in Japan.

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 or krill, Decapoda (shrimp, lobsters, crabs), and the minuscule Amphionides.

The order Euphasiacea comprises two families. The more abundant Euphausiidae contains ten different genera with a total of 85 species. Of these, the genus Euphausia is the largest, with 31 species.^[2] The lesser known family, the Bentheuphausiidae, has only one species, Bentheuphausia amblyops, a bathypelagic krill living in deep waters below 1,000 metres (3,300 ft). It is considered the most primitive living species of all krill.^[3]

Well-known species of the Euphausiidae of commercial krill fisheries include Antarctic krill (Euphausia superba), Pacific krill (Euphausia pacifica) and Northern krill (Meganyctiphanes norvegica).^[4]

Phylogeny

Main article: Phylogeny of Malacostraca

See also: Eucarida#phylogeny

Proposed phylogeny of Euphausiacea[5]

Euphausiacea Bentheuphausiidae Bentheuphausia Euphausiidae Thysanopoda (♣) Nematobrachion (♦) Euphausiinae Meganyctiphanes Euphausiini (♠)(♦) Pseudeuphausia Euphausia Nematoscelini (♠) Nyctiphanes Nematoscelina (♠) Nematoscelis Thysanoessa Tessarabrachion Stylocheiron

Phylogeny obtained from morphological data, (♠) names coined in,[5] (♣) possibly paraphyletic taxon due to Nematobrachion in.[5] (♦) clades differs from Casanova (1984),[6] where Pseudoeuphausia is sister to Nyctiphanes, Euphausia is sister to Thysanopoda and Nematobrachion is sister to Stylocheiron.

The order Euphausiacea is believed to be monophyletic due to several unique conserved morphological characteristics (autapomorphy) such as the naked filamentous gills or the thin thoracopods.^[7] and by molecular studies.^[8][9]

There have been many theories of the location of the order Euphausiacea, in fact 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 of Schizopoda, which was split by Johan Erik Vesti Boas in 1883 into two separate orders.^[10] Later, William Thomas Calman (1904) ranked the Mysidacea in the Peracarida super-order and euphausiids in Eucarida super-order, although even up to the 1930s the order Schizopoda was advocated.^[7] 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^[11] and Isabella Gordon.^[12] The reason for this debate is that krill share some morphological features of decapods and others of mysids.^[7]

Molecular studies have also no been able to unambiguously group them, possibly due to the lack of many key rare species such as Bentheuphausia amblyops in krill and Amphionides reynaudii in Eucarida. One study supports Eucarida monophyly (with basal Mysida),^[13] another groups Euphausiacea with Mysida (the Schizopoda),^[9] while yet another groups Euphausiacea with Hoplocarida.^[14]

Timeline

Unusual for crustaceans, no fossil has been found that can be unequivocally assigned to the order Euphausiacea. Some extinct eumalacostracan taxa have been thought to be euphausiaceans such as Anthracophausia, Crangopsis —now assigned to the Aeschronectida (Hoplocarida)^[5]—and Palaeomysis.^[15] Consequently the dating of the speciation events have been estimated by means of molecular clock methods, which place 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.^[9]

Distribution

A krill swarm

Krill occur worldwide in all oceans, although many individual species have endemic or neritic (i.e., coastal) restricted distributions. Bentheuphausia amblyops, a bathypelagic species, has a cosmopolitan distribution within its deep-sea habitat.^[16]

Species of the genus Thysanoessa occur in both the Atlantic and Pacific oceans.^[17] The Pacific is home to Euphausia pacifica. Northern krill occur across the Atlantic from the Mediterranean Sea northward.

Species with neritic distributions include the four species of the genus Nyctiphanes.^[8] They are highly abundant along the upwelling regions of the California, Humboldt, Benguela, and Canarias current systems.^[17][18][19] Another species having only neritic distribution is E. crystallorophias, which occurs only along the Antarctic coastline (and thus also is endemic to that region).^[20]

Species with endemic distributions include Nyctiphanes capensis, which occurs only in the Benguela current,^[8] E. mucronata in the Humboldt current,^[21] and the six Euphausia species native to the Southern Ocean.

In the Antarctic, seven species are known,^[22] one species of the genus Thysanoessa (T. macrura) and six of the genus Euphausia. The Antarctic krill (Euphausia superba) commonly lives at depths of as much as 100 m (330 ft),^[23] whereas ice krill (Euphausia crystallorophias) have been recorded at a depth of 4,000 m (13,100 ft), though they commonly live at depths of at most 300–600 m (1,000–2,000 ft).^[24] Both are found at latitudes south of 55° S, with E. crystallorophias dominating south of 74° S^[25] and in regions of pack ice. Other species known in the Southern Ocean are E. frigida, E. longirostris, E. triacantha and E. vallentini.^[26]

Anatomy and morphology

Krill anatomy explained, using Euphausia superba as a model

Krill are crustaceans and have a chitinous exoskeleton made up of three segments: the cephalon (head), the thorax, and the abdomen. The first two segments are fused into one segment, the cephalothorax. This outer shell of krill is transparent in most species. Krill feature intricate compound eyes; some species can adapt to different lighting conditions through the use of screening pigments.^[27] 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 the feeding legs and the grooming legs. Additionally all species have five 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.^[28] Krill can be easily distinguished from other crustaceans such as true shrimp by their externally visible gills.^[29]

The gills of krill are externally visible.

Many krill are filter feeders:^[18] their frontmost appendages, the thoracopods, form very fine combs with which they can filter out their food from the water. These 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. However, it is believed that krill are mostly omnivorous.^[30] A few species are carnivorous, preying on small zooplankton and fish larvae.^[31]

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^[32] and that the krill probably do not produce this substance themselves but acquire it as part of their diet, which contains dinoflagellates.^[33] Krill photophores are complex organs with lenses and focusing abilities, and they can be rotated by muscles.^[34] The precise function of these organs is as yet unknown; they might have a purpose in mating, social interaction or orientation. Some researchers (e.g., Lindsay & Latz and Johnsen) have proposed that krill use the light as a form of counter-illumination camouflage to compensate their shadow against the ambient light from above to make themselves less visible to predators from below.^[35][36]

Behaviour

Most krill are swarming animals; the sizes and densities of such swarms vary greatly depending on the species and the region. For Euphausia superba, there have been reports of swarms of up to 10,000 to 60,000 individuals per cubic metre.^[37][38] Swarming is a defensive mechanism, confusing smaller predators that would like to pick out single individuals. Krill typically follow a diurnal vertical migration. Until recently it has been assumed that they spend the day at greater depths and rise during the night toward the surface. It has been found that the deeper they go, the more they reduce their activity,^[39] apparently to reduce encounters with predators and to conserve energy. Later work suggested that swimming activity in krill varied with stomach fullness. Satiated animals that had been feeding at the surface swim less actively and therefor sink below the mixed layer.^[40] As they sink they produce faeces which may mean that they have an important role to play in the Antarctic carbon cycle. Krill with empty stomachs were found to swim more actively and thus head towards the surface. This implies that vertical migration may be a bi or tri daily occurrence. Some species (e.g., Euphausia superba, E. pacifica, E. hanseni, Pseudeuphausia latifrons, and Thysanoessa spinifera) also form surface swarms during the day for feeding and reproductive purposes even though such behaviour is dangerous because it makes them extremely vulnerable to predators.^[41]

Beating pleopods of a swimming Antarctic krill

Dense swarms may elicit a feeding frenzy among fish, birds and mammal predators, especially near the surface. When disturbed, a swarm scatters, and some individuals have even been observed to moult instantaneously, leaving the exuvia behind as a decoy.^[42]

Krill normally swim at pace of 5–10 cm/s (2–3 body lengths per second),^[43] using their swimmerets for propulsion. Their larger migrations are subject to the currents in the ocean. 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).^[44] Their swimming performance has led many researchers to classify adult krill as micro-nektonic life-forms, i.e., small animals capable of individual motion against (weak) currents. Larval forms of krill are generally considered zooplankton.^[4]

Ecology and life history

See also: Carbon sequestration and biological pump

NASA SeaWiFS satellite image of the large phytoplankton bloom in the Bering Sea in 1998

Krill are an important element of the food chain. Antarctic krill feed directly on phytoplankton, converting the primary production energy into a form suitable for consumption by larger animals that cannot feed directly on the minuscule algae. Some species like the Northern krill have a relatively small filtering basket and actively hunt for copepods and larger zooplankton.^[31] Many animals feed on krill, ranging from smaller animals like fish or penguins to larger ones like seals and even baleen whales.^[45]

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,^[46] 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, and the incident was even thought to have been a reason for salmon not returning to the rivers of western Alaska that season.^[47]

Other factors besides predation and food availability can influence the mortality rate in krill populations. As temperatures have risen over the past couple decades, Antarctic sea ice has melted. In this way, climate change poses a threat to krill populations as they feed on algae beneath the ice.^[48] There are several single-celled endoparasitoidic ciliates of the genus Collinia that can infect different species of krill and cause massive decline in affected populations. Such diseases have been reported for Thysanoessa inermis in the Bering Sea and also for E. pacifica, Thysanoessa spinifera, and T. gregaria off the North American Pacific coast.^[49][50] There are also some ectoparasites of the family Dajidae (epicaridean isopods) that afflict krill (and also shrimp and mysids); one such parasite is Oculophryxus bicaulis, which has been found on the krill Stylocheiron affine and S. longicorne. It attaches itself to the eyestalk of the animal and sucks blood from its head; it is believed that it inhibits the reproduction of its host, as none of the afflicted animals found reached maturity.^[51]

Life history

A nauplius of Euphausia pacifica hatching, emerging backwards from the egg

The general life cycle of krill has been the subject of several studies (e.g., Gurney, 1942^[11] and Mauchline & Fisher, 1969^[18]) performed on a variety of species and is thus relatively well understood, although there are minor variations in detail from species to species. After krill hatch from the egg, they go through several larval stages called the nauplius, pseudometanauplius, metanauplius, calyptopsis, and furcilia stages, each of which is sub-divided into several 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 multiple times as they develop, shedding their rigid exoskeleton whenever it becomes too small and growing a new one. Smaller animals moult more frequently than larger ones. Up through the metanauplius stage, the larvae are nourished by yolk reserves within their body. Only by the calyptopsis stages has differentiation progressed far enough for them to develop a mouth and a digestive tract, and they begin to feed upon phytoplankton. By that time, the larvae must have reached the photic zone, the upper layers of the ocean where algae flourish, for their yolk reserves are exhausted by then and they would starve otherwise. 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.^[52] After the final furcilia stage, the krill emerges in a shape similar to an adult, but it is still an immature juvenile, that only subsequently develops gonads and matures.^[53]

During the mating season, which varies depending on the species and the climate, the male deposits a sperm sack at the genital opening (named thelycum) of the female. 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.^[54] Krill can have multiple broods in one season, with interbrood periods in the order of days.^[19][55]

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 millimetre (0.012–0.016 in).

There are two types of spawning mechanism.^[19] 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 into deeper waters, 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.^[56] 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 difficilis may hatch as nauplius or pseudometanauplius.^[57]

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.^[4] Subtropical or tropical species' longevity is still shorter, e.g., Nyctiphanes simplex, which usually lives for only six to eight months.^[58]

Moulting occurs whenever the animal outgrows its rigid exoskeleton. Young animals, growing faster, moult more often than older and larger ones. The frequency of moulting varies widely from species to species and is, even within one species, subject to many external factors such as the latitude, the water temperature, and the availability of food. The subtropical species Nyctiphanes simplex, for instance, has an overall inter-moult period in the range 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 (37 and 59 °F).^[59] E. superba is able to reduce its body size when there is not enough food available, moulting also when its exoskeleton becomes too large.^[60] 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 abnormally high water temperatures. Shrinkage has been postulated for other temperate-zone species of krill as well.^[61]

Economy

See also: Krill fishery

Deep frozen plates of Antarctic krill for use as animal feed and raw material for cooking

Krill has been harvested as a food source for humans (okiami) and domesticated animals since the 19th century, in Japan maybe even earlier. Large-scale fishing developed only 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. A peak in krill harvest had been reached in 1983 with more than 528,000 tonnes in the Southern Ocean alone (of which the Soviet Union produced 93%). In 1993, two events led to a drastic decline in krill production: first, Russia abandoned its operations, and second, the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) defined maximum catch quotas for a sustainable exploitation of Antarctic krill. The annual catch in Antarctic waters seems to have stabilised around 100,000 tonnes of krill, which is roughly one fiftieth of the CCAMLR catch quota.^[62] The main limiting factor is probably the high cost associated with Antarctic operations, although there are some political and legal issues as well.^[63] The fishery around Japan appears to have saturated at some 70,000 tonnes.^[64]

Experimental small-scale harvesting is being carried out in other areas, for example, fishing for Euphausia pacifica off British Columbia and harvesting Meganyctiphanes norvegica, Thysanoessa raschii and Thysanoessa inermis in the Gulf of St. Lawrence. These experimental operations produce only a few hundred tonnes of krill per year. Nicol & Foster consider it unlikely that any large-scale harvesting operations in these areas will be started due to opposition from local fishing industries and conservation groups.^[64]

Krill tastes salty and somewhat stronger than shrimp. For mass-consumption and commercially prepared products they must be peeled, because their exoskeleton contains fluorides, which are toxic in high concentrations.^[65] There is a small but growing market for krill oil as a dietary supplement ingredient. Two clinical trials have been published; tests included lipid lowering, arthritis pain and function, and C-reactive protein.^[66][67]

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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 the Marine Biological Association of the United Kingdom No. 15, Plymouth, UK, 2003.
• Everson, I. (ed.): Krill: biology, ecology and fisheries. Oxford, Blackwell Science; 2000. ISBN 0-632-05565-0.
• 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 Biology from Napier University.
• Tett, P.: Bioluminescence, lecture notes from the 1999/2000 edition of that same course.

External links

Crustaceans portal

Media related to Krill at Wikimedia Commons Data related to Euphausia at Wikispecies

• Webcam of Krill Aquarium at Australian Antarctic Division
• The Wiktionary definition of krill
• 'Antarctic Energies' animation by Lisa Roberts