Brine shrimp

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Brine shrimp
Artemia salina 4.jpg
Artemia salina mating pair – female left, male right
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Branchiopoda
Subclass: Sarsostraca
Order: Anostraca
Suborder: Artemiina
Family: Artemiidae
Grochowski, 1895
Genus: Artemia
Leach, 1819
Species [1]

Artemia is a genus of aquatic crustaceans also known as brine shrimp or sea monkeys. It is the only genus in the family Artemiidae. The first historical record of the existence of Artemia dates back to the first half of the 10th century AD from Lake Urmia, Iran, with an example called by an Iranian geographer an "aquatic dog", [2] although the first unambiguous record is the report and drawings made by Schlösser in 1757 of animals from Lymington, England. [3] Artemia populations are found worldwide, typically in inland saltwater lakes, but occasionally in oceans. Artemia are able to avoid cohabiting with most types of predators, such as fish, by their ability to live in waters of very high salinity (up to 25%). [4]

Contents

The ability of the Artemia to produce dormant eggs, known as cysts, has led to extensive use of Artemia in aquaculture. The cysts may be stored indefinitely and hatched on demand to provide a convenient form of live feed for larval fish and crustaceans. [4] Nauplii of the brine shrimp Artemia constitute the most widely used food item, and over 2,000 metric tons (2,200 short tons) of dry Artemia cysts are marketed worldwide annually with most of the cysts being harvested from the Great Salt Lake in Utah. [5] In addition, the resilience of Artemia makes them ideal animals for running biological toxicity assays and it has become a model organism used to test the toxicity of chemicals. Breeds of Artemia are sold as novelty gifts under the marketing name Sea-Monkeys .

Description

The brine shrimp Artemia comprises a group of seven to nine species very likely to have diverged from an ancestral form living in the Mediterranean area about 5.5  million years ago, [6] around the time of the Messinian salinity crisis.

The Laboratory of Aquaculture & Artemia Reference Center at Ghent University possesses the largest known Artemia cyst collection, a cyst bank containing over 1,700 Artemia population samples collected from different locations around the world. [7]

Artemia is a typical primitive arthropod with a segmented body to which is attached broad leaf-like appendages. The body usually consists of 19 segments, the first 11 of which have pairs of appendages, the next two which are often fused together carry the reproductive organs, and the last segments lead to the tail. [8] The total length is usually about 8–10 millimetres (0.31–0.39 in) for the adult male and 10–12 mm (0.39–0.47 in) for the female, but the width of both sexes, including the legs, is about 4 mm (0.16 in).

The body of Artemia is divided into head, thorax, and abdomen. The entire body is covered with a thin, flexible exoskeleton of chitin to which muscles are attached internally and which is shed periodically. [9] In female Artemia, a moult precedes every ovulation.

For brine shrimp, many functions, including swimming, digestion and reproduction are not controlled through the brain; instead, local nervous system ganglia may control some regulation or synchronisation of these functions. [9] Autotomy, the voluntary shedding or dropping of parts of the body for defence, is also controlled locally along the nervous system. [8] Artemia have two types of eyes. They have two widely separated compound eyes mounted on flexible stalks. These compound eyes are the main optical sense organ in adult brine shrimps. The median eye, or the naupliar eye, is situated anteriorly in the centre of the head and is the only functional optical sense organ in the nauplii, which is functional until the adult stage. [9]

Ecology and behavior

Brine shrimp can tolerate any levels of salinity from 25‰ to 250‰ (25–250 g/L), [10] with an optimal range of 60‰–100‰, [10] and occupy the ecological niche that can protect them from predators. [11] Physiologically, optimal levels of salinity are about 30–35‰, but due to predators at these salt levels, brine shrimp seldom occur in natural habitats at salinities of less than 60–80‰. Locomotion is achieved by the rhythmic beating of the appendages acting in pairs. Respiration occurs on the surface of the legs through fibrous, feather-like plates (lamellar epipodites). [8]

An Artemia cyst Brine shrimp cyst.jpg
An Artemia cyst

Reproduction

Males differ from females by having the second antennae markedly enlarged, and modified into clasping organs used in mating. [12] Adult female brine shrimp ovulate approximately every 140 hours. In favourable conditions, the female brine shrimp can produce eggs that almost immediately hatch.[ citation needed ] While in extreme conditions, such as low oxygen level or salinity above 150‰, female brine shrimp produce eggs with a chorion coating which has a brown colour. These eggs, also known as cysts, are metabolically inactive and can remain in total stasis for two years while in dry oxygen-free conditions, even at temperatures below freezing. This characteristic is called cryptobiosis, meaning "hidden life". While in cryptobiosis, brine shrimp eggs can survive temperatures of liquid air (−190 °C or −310 °F) and a small percentage can survive above boiling temperature (105 °C or 221 °F) for up to two hours. [11] Once placed in briny (salt) water, the eggs hatch within a few hours. The nauplius larvae are less than 0.4 mm in length when they first hatch.

Parthenogenesis

The effects of central fusion and terminal fusion on heterozygosity Central fusion and terminal fusion automixis.svg
The effects of central fusion and terminal fusion on heterozygosity

Parthenogenesis is a natural form of reproduction in which growth and development of embryos occur without fertilisation. Thelytoky is a particular form of parthenogenesis in which the development of a female individual occurs from an unfertilised egg. Automixis is a form of thelytoky, but there are different kinds of automixis. The kind of automixis relevant here is one in which two haploid products from the same meiosis combine to form a diploid zygote.

Diploid Artemia parthenogenetica reproduce by automictic parthenogenesis with central fusion (see diagram) and low but nonzero recombination. [13] Central fusion of two of the haploid products of meiosis (see diagram) tends to maintain heterozygosity in transmission of the genome from mother to offspring, and to minimise inbreeding depression. Low crossover recombination during meiosis likely restrains the transition from heterozygosity to homozygosity over successive generations.

Diet

In their first stage of development, Artemia do not feed but consume their own energy reserves stored in the cyst. [14] Wild brine shrimp eat microscopic planktonic algae. Cultured brine shrimp can also be fed particulate foods including yeast, wheat flour, soybean powder or egg yolk. [15]

Genetics, genomics and transcriptomics

Artemia comprises sexually reproducing, diploid species and several obligate parthenogenetic Artemia populations consisting of different clones and ploidies (2n->5n). [16] Several genetic maps have been published for Artemia. [17] [18] The past years, different transcriptomic studies have been performed to elucidate biological responses in Artemia, such as its response to salt stress, [19] [20] toxins, [21] infection [22] and diapause termination. [23] These studies also led to various fully assembled Artemia transcriptomes. Recently, the Artemia genome was assembled and annotated, revealing a genome containing an unequaled 58% of repeats, genes with unusually long introns and adaptations unique to the extremophilic nature of Artemia in high salt and low oxygen environments. [24] These adaptations include a unique energy-intensive endocytosis-based salt excretion strategy resembling salt excretion strategies of plants, as well as several survival strategies for extreme environments it has in common with the extremophilic tardigrade. [24]

Aquaculture

San Francisco Bay Salt Ponds Artemia breeding ponds.jpg
San Francisco Bay Salt Ponds

Fish farm owners search for a cost-effective, easy to use, and available food that is preferred by the fish. From cysts, brine shrimp nauplii can readily be used to feed fish and crustacean larvae just after a one-day incubation. Instar I (the nauplii that just hatched and with large yolk reserves in their body) and instar II nauplii (the nauplii after first moult and with functional digestive tracts) are more widely used in aquaculture, because they are easy for operation, rich in nutrients, and small, which makes them suitable for feeding fish and crustacean larvae live or after drying.

Toxicity test

Artemia found favor as a model organism for use in toxicological assays, despite the recognition that it is too robust an organism to be a sensitive indicator species. [25]

In pollution research Artemia, the brine shrimp, has had extensive use as a test organism and in some circumstances is an acceptable alternative to the toxicity testing of mammals in the laboratory. [26] The fact that millions of brine shrimp are so easily reared has been an important help in assessing the effects of a large number of environmental pollutants on the shrimps under well controlled experimental conditions.

Conservation

Artemia monica (male) Artemia monica.jpg
Artemia monica (male)

Overall, brine shrimp are abundant, but some populations and localized species do face threats, especially from habitat loss to introduced species. For example, A. franciscana of the Americas has been widely introduced to places outside its native range and is often able to outcompete local species, such as A. salina in the Mediterranean region. [27] [28]

Among the highly localized species are A. urmiana from Lake Urmia in Iran. Once abundant, the species has drastically declined due to drought, leading to fears that it was almost extinct. [29] However, a second population of this species has recently been discovered in the Koyashskoye Salt Lake at the Crimean Peninsula. [30]

A. monica , the species commonly known as Mono Lake brine shrimp, can be found in Mono Lake, Mono County, California. In 1987, Dennis D. Murphy from Stanford University petitioned the United States Fish and Wildlife Service to add A. monica to the endangered species list under the Endangered Species Act (1973). The diversion of water by the Los Angeles Department of Water and Power resulted in rising salinity and concentration of sodium hydroxide in Mono Lake. Despite the presence of trillions of brine shrimp in the lake, the petition contended that the increase in pH would endanger them. The threat to the lake's water levels was addressed by a revision to California State Water Resources Control Board's policy, and the US Fish and Wildlife Service found on 7 September 1995 that the Mono Lake brine shrimp did not warrant listing. [31]

Space experiment

Scientists have taken the eggs of brine shrimp to outer space to test the impact of radiation on life. Brine shrimp cysts were flown on the U.S. Biosatellite 2, Apollo 16, and Apollo 17 missions, and on the Russian Bion-3 (Cosmos 782), Bion-5 (Cosmos 1129), Foton 10, and Foton 11 flights. Some of the Russian flights carried European Space Agency experiments.

On Apollo 16 and Apollo 17, the cysts traveled to the Moon and back. Cosmic rays that passed through an egg would be detected on the photographic film in its container. Some eggs were kept on Earth as experimental controls as part of the tests. Also, as the take-off in a spacecraft involves a lot of shaking and acceleration, one control group of egg cysts was accelerated to seven times the force of gravity and vibrated mechanically from side to side for several minutes so that they could experience the same violence of a rocket take-off. [32] About 400 eggs were in each experimental group. All the egg cysts from the experiment were then placed in salt water to hatch under optimum conditions. The results showed A. salina eggs are highly sensitive to cosmic radiation; 90% of the embryos induced to develop from hit eggs died at different developmental stages. [33]

Related Research Articles

A halophile is an extremophile that thrives in high salt concentrations. In chemical terms, halophile refers to a Lewis acidic species that has some ability to extract halides from other chemical species.

<span class="mw-page-title-main">Mono Lake</span> Endorheic lake in California

Mono Lake is a saline soda lake in Mono County, California, formed at least 760,000 years ago as a terminal lake in an endorheic basin. The lack of an outlet causes high levels of salts to accumulate in the lake which make its water alkaline.

<span class="mw-page-title-main">Great Salt Lake</span> Salt lake in Utah, United States

The Great Salt Lake is the largest saltwater lake in the Western Hemisphere and the eighth-largest terminal lake in the world. It lies in the northern part of the U.S. state of Utah and has a substantial impact upon the local climate, particularly through lake-effect snow. It is a remnant of Lake Bonneville, a prehistoric body of water that covered much of western Utah.

<span class="mw-page-title-main">Black-necked grebe</span> Water bird from parts of Africa, Eurasia, and the Americas

The black-necked grebe or eared grebe is a member of the grebe family of water birds. It was described in 1831 by Christian Ludwig Brehm. There are currently three accepted subspecies, including the nominate subspecies. Its breeding plumage features distinctive ochre-coloured feathers which extend behind its eye and over its ear coverts. The rest of the upper parts, including the head, neck, and breast, are coloured black to blackish brown. The flanks are tawny rufous to maroon-chestnut, and the abdomen is white. In its non-breeding plumage, this bird has greyish-black upper parts, including the top of the head and a vertical stripe on the back of the neck. The flanks are also greyish-black. The rest of the body is a white or whitish colour. The juvenile has more brown in its darker areas. The subspecies californicus can be distinguished from the nominate by the former's usually longer bill. The other subspecies, P. n. gurneyi, can be differentiated by its greyer head and upper parts and by its smaller size. P. n. gurneyi can also be told apart by its lack of a non-breeding plumage. This species is present in parts of Africa, Eurasia, and the Americas.

Sea-Monkeys is a marketing term for brine shrimp (Artemia) sold as novelty aquarium pets. Developed in the United States in 1957 by Harold von Braunhut, they are sold as eggs intended to be added to water, and most often come bundled in a kit of three pouches and instructions. Sometimes a small tank and additional pouches are included. The product was marketed in the 1960s and 70s, especially in comic books, and remains a presence in popular culture.

<span class="mw-page-title-main">Mysida</span> Small, shrimp-like crustacean

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 pouch and are not free-swimming characterises the order. The mysid's head bears a pair of stalked eyes and two pairs of antennae. The thorax consists of eight segments each bearing branching limbs, the whole concealed beneath a protective carapace and the abdomen has six segments and usually further small limbs.

<span class="mw-page-title-main">Salt lake</span> Landlocked body of water which has a high concentration of salts

A salt lake or saline lake is a landlocked body of water that has a concentration of salts and other dissolved minerals significantly higher than most lakes. In some cases, salt lakes have a higher concentration of salt than sea water; such lakes can also be termed hypersaline lakes, and may also be pink lakes on account of their colour. An alkalic salt lake that has a high content of carbonate is sometimes termed a soda lake.

<span class="mw-page-title-main">Lake Urmia</span> Salt lake in Iran

Lake Urmia is an endorheic salt lake in Iran. The lake is located between the provinces of East Azerbaijan and West Azerbaijan in Iran, and west of the southern portion of the Caspian Sea. At its greatest extent, it was the largest lake in the Middle East and the sixth-largest saltwater lake on Earth, with a surface area of approximately 5,200 km2 (2,000 sq mi), a length of 140 km (87 mi), a width of 55 km (34 mi), and a maximum depth of 16 m (52 ft).

<span class="mw-page-title-main">Anostraca</span> Order of crustaceans

Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered mountain lakes, and Antarctic ice. They are usually 6–25 mm (0.24–0.98 in) long. Most species have 20 body segments, bearing 11 pairs of leaf-like phyllopodia, and the body lacks a carapace. They swim "upside-down" and feed by filtering organic particles from the water or by scraping algae from surfaces, with the exception of Branchinecta gigas, or "giant fairy shrimp", which is itself a predator of other species of anostracans. They are an important food for many birds and fish, and some are cultured and harvested for use as fish food. There are 300 species spread across 8 families.

<span class="mw-page-title-main">Cryptobiosis</span> Metabolic state of life

Cryptobiosis or anabiosis is a metabolic state in extremophilic organisms in response to adverse environmental conditions such as desiccation, freezing, and oxygen deficiency. In the cryptobiotic state, all measurable metabolic processes stop, preventing reproduction, development, and repair. When environmental conditions return to being hospitable, the organism will return to its metabolic state of life as it was prior to cryptobiosis.

<span class="mw-page-title-main">Thelytoky</span> Type of parthenogenesis in which females are produced from unfertilized eggs

Thelytoky is a type of parthenogenesis and is the absence of mating and subsequent production of all female diploid offspring as for example in aphids. Thelytokous parthenogenesis is rare among animals and reported in about 1,500 species, about 1 in 1000 of described animal species, according to a 1984 study. It is more common in invertebrates, like arthropods, but it can occur in vertebrates, including salamanders, fish, and reptiles such as some whiptail lizards.

<i>Dunaliella</i> Genus of algae

Dunaliella is a single-celled, photosynthetic green alga, that is characteristic for its ability to outcompete other organisms and thrive in hypersaline environments. It is mostly a marine organism, though there are a few freshwater species that tend to be more rare. It is a genus in which certain species can accumulate relatively large amounts of β-carotenoids and glycerol in very harsh growth conditions consisting of high light intensities, high salt concentrations, and limited oxygen and nitrogen levels, yet is still very abundant in lakes and lagoons all around the world.

<i>Artemia salina</i> Species of small brine shrimp

Artemia salina is a species of brine shrimp – aquatic crustaceans that are more closely related to Triops and cladocerans than to true shrimp. It belongs to a lineage that does not appear to have changed much in 100 million years.

<span class="mw-page-title-main">Aquaculture of brine shrimp</span> Farming of a type of crustacean

Brine shrimp have the ability to produce dormant eggs, known as cysts. This has led to the extensive use of brine shrimp in aquaculture. The cysts may be stored for long periods and hatched on demand to provide a convenient form of live feed for larval fish and crustaceans.

<i>Tigriopus brevicornis</i> Coastal marine copepod from north western Europe

Tigriopus brevicornis is a coastal marine copepod. They are a dominant member of shallow supra tidal rock pools along the North Western European coastline. A broad range of studies have been carried out on this species, including: its ecology, physiology, phylogeography, metapopulation genetics, development and reproductive behaviour. T. brevicornis has also recently been used in ecotoxicology studies and has been trialled as a live feed for larvae in several aquaculture-based studies for the past 30 years.

<i>Artemia monica</i> Species of small freshwater animal

Artemia monica, the Mono Lake brine shrimp, is a species of brine shrimp, endemic to Mono Lake in California, United States.

<i>Artemia parthenogenetica</i> Species of small freshwater animal

Artemia parthenogenetica is a species of brine shrimp – aquatic crustaceans belonging to a different class, the Branchiopoda, than the true shrimps.

Flamingolepis liguloides is a parasitic tapeworm of the Cestoda class. There are several tapeworms that are found to infect Artemia; however, F. liguloides is the most prevalent species of infectious tapeworm among Artemia. F. liguloides infects brine shrimp (Artemia) as the intermediate host and flamingos as the definitive host. Effects of the tapeworm in flamingos is unclear, though researchers hypothesize that a high parasitemia could potentially be deadly to the host. The parasite appears to affect the Artemia spp. as it alters the behavior and color of its host.

James Standish Clegg is a Professor Emeritus of Biochemistry at University of California, Davis, based at the Coastal and Marine Sciences Institute (CMSI) in Bodega Bay, California. He served as director of the Bodega Marine Laboratory (BML) from 1986 to 1999 and as president of the National Association Marine Laboratories (NAML) from 1991 to 1993.

<i>Artemia franciscana</i> Species of crustacean

Artemia franciscana is a species of brine shrimp endemic to the Americas but now widely introduced throughout the tropics and temperate zones worldwide.

References

  1. ↑ Alireza Asem (2023). "Phylogenetic analysis of problematic Asian species of Artemia Leach, 1819 (Crustacea, Anostraca), with the descriptions of two new species". Journal of Crustacean Biology. 83: 1–25.
  2. ↑ Alireza Asem; Amin Eimanifar (2016). "Updating historical record on brine shrimp Artemia (Crustacea: Anostraca) from Urmia Lake (Iran) in the first half of the 10th century AD" (PDF). International Journal of Aquatic Science. 7: 1–5. Archived from the original (PDF) on 2016-04-01. Retrieved 2016-11-24.
  3. ↑ Alireza Asem (2008). "Historical record on brine shrimp Artemia more than one thousand years ago from Urmia Lake, Iran" (PDF). Journal of Biological Research-Thessaloniki. 9: 113–114. Archived from the original (PDF) on 2016-12-01. Retrieved 2013-05-17.
  4. 1 2 Martin Daintith (1996). Rotifers and Artemia for Marine Aquaculture: a Training Guide. University of Tasmania. OCLC   222006176.
  5. ↑ "Introduction, biology and ecology of Artemia" . Retrieved 15 October 2022.
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  10. 1 2 John K. Warren (2006). "Halotolerant life in feast or famine (a source of hydrocarbons and a fixer of metals)". Evaporites: Sediments, Resources and Hydrocarbons. Birkhäuser. pp.  617–704. ISBN   978-3-540-26011-0.
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  15. ↑ Kai Schumann (August 10, 1997). "Artemia (Brine Shrimp) FAQ 1.1". Portland State University. Archived from the original on August 14, 2007. Retrieved March 13, 2010.
  16. ↑ Maniatsi, Stefania; Baxevanis, Athanasios D.; Kappas, Ilias; Deligiannidis, Panagiotis; Triantafyllidis, Alexander; Papakostas, Spiros; Bougiouklis, Dimitrios; Abatzopoulos, Theodore J. (2011-02-01). "Is polyploidy a persevering accident or an adaptive evolutionary pattern? The case of the brine shrimp Artemia". Molecular Phylogenetics and Evolution. 58 (2): 353–364. doi: 10.1016/j.ympev.2010.11.029 . PMID   21145977.
  17. ↑ Vos, Stephanie De; Bossier, Peter; Stappen, Gilbert Van; Vercauteren, Ilse; Sorgeloos, Patrick; Vuylsteke, Marnik (2013-03-04). "A first AFLP-Based Genetic Linkage Map for Brine Shrimp Artemia franciscana and Its Application in Mapping the Sex Locus". PLOS ONE. 8 (3): e57585. Bibcode:2013PLoSO...857585D. doi: 10.1371/journal.pone.0057585 . ISSN   1932-6203. PMC   3587612 . PMID   23469207.
  18. ↑ Han, Xuekai; Ren, Yizhuo; Ouyang, Xuemei; Zhang, Bo; Sui, Liying (2021-07-15). "Construction of a high-density genetic linkage map and QTL mapping for sex and growth traits in Artemia franciscana". Aquaculture. 540: 736692. Bibcode:2021Aquac.54036692H. doi:10.1016/j.aquaculture.2021.736692. ISSN   0044-8486. S2CID   233663524.
  19. ↑ De Vos, S.; Van Stappen, G.; Sorgeloos, P.; Vuylsteke, M.; Rombauts, S.; Bossier, P. (2019-02-01). "Identification of salt stress response genes using the Artemia transcriptome". Aquaculture. 500: 305–314. Bibcode:2019Aquac.500..305D. doi:10.1016/j.aquaculture.2018.09.067. ISSN   0044-8486. S2CID   92842322.
  20. ↑ Lee, Junmo; Park, Jong Soo (2020-10-13). "Tolerance at the genetic level of the brine shrimp Artemia salina to a wide range of salinity". www.researchsquare.com. doi:10.21203/rs.3.rs-91049/v1. S2CID   234621039 . Retrieved 2021-09-02.
  21. ↑ Yi, Xianliang; Zhang, Keke; Liu, Renyan; Giesy, John P.; Li, Zhaochuan; Li, Wentao; Zhan, Jingjing; Liu, Lifen; Gong, Yufeng (2020-01-01). "Transcriptomic responses of Artemia salina exposed to an environmentally relevant dose of Alexandrium minutum cells or Gonyautoxin2/3". Chemosphere. 238: 124661. Bibcode:2020Chmsp.23824661Y. doi:10.1016/j.chemosphere.2019.124661. ISSN   0045-6535. PMID   31472350. S2CID   201700530.
  22. ↑ Zhang, Yulong; Wang, Di; Zhang, Zao; Wang, Zhangping; Zhang, Daochuan; Yin, Hong (2018-10-01). "Transcriptome analysis of Artemia sinica in response to Micrococcus lysodeikticus infection". Fish & Shellfish Immunology. 81: 92–98. doi:10.1016/j.fsi.2018.06.033. ISSN   1050-4648. PMID   30006042. S2CID   51624497.
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  24. 1 2 De Vos, Stephanie; Rombauts, Stephane; Coussement, Louis; Dermauw, Wannes; Vuylsteke, Marnik; Sorgeloos, Patrick; Clegg, James S.; Nambu, Ziro; Van Nieuwerburgh, Filip; Norouzitallab, Parisa; Van Leeuwen, Thomas (2021-08-31). "The genome of the extremophile Artemia provides insight into strategies to cope with extreme environments". BMC Genomics. 22 (1): 635. doi: 10.1186/s12864-021-07937-z . ISSN   1471-2164. PMC   8406910 . PMID   34465293.
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  28. ↑ Hachem Ben Naceur; Amel Ben Rejeb Jenhani; Mohamed Salah Romdhane (2009). "New distribution record of the brine shrimp Artemia (Crustacea, Branchiopoda, Anostraca) in Tunisia". Check List. 5 (2): 281–288. doi: 10.15560/5.2.281 . ISSN   1809-127X.
  29. ↑ "Lake Urumia's Artemia Face Extinction". Financial Tribune. 28 December 2014. Retrieved 29 January 2018.
  30. ↑ Eimanifar A; Asem A; Djamali M; Wink M (2016). "A note on the biogeographical origin of the brine shrimp Artemia urmiana Günther, 1899 from Urmia Lake, Iran" (PDF). Zootaxa. 4097 (2): 294–300. doi:10.11646/zootaxa.4097.2.12. PMID   27394547. S2CID   25998873. Archived from the original (PDF) on 2020-02-10.
  31. ↑ "Endangered and Threatened Wildlife and Plants; 12-Month Finding for a Petition to List the Mono Lake Brine Shrimp as Endangered". Federal Register. 60 (173): 46571–46572. 1995.[ permanent dead link ]
  32. ↑ H. Planel; Y. Gaubin; R. Kaiser; B. Pianezzi (1980). "The effects of cosmic rays on Artemia egg cysts". Laboratoire Médicale. Report for National Aeronautics and Space Administration.
  33. ↑ H. Bücker & G. Horneck (1975). "The biological effectiveness of HZE-particles of cosmic radiation studied in the Apollo 16 and 17 Biostack experiments". Acta Astronautica . 2 (3–4): 247–264. Bibcode:1975AcAau...2..247B. doi:10.1016/0094-5765(75)90095-8. PMID   11887916.
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