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Datasheet

Artemia
(brine shrimp)

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Datasheet

Artemia (brine shrimp)

Summary

  • Last modified
  • 14 July 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Animal
  • Preferred Scientific Name
  • Artemia
  • Preferred Common Name
  • brine shrimp
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Crustacea
  •         Class: Branchiopoda
  • Summary of Invasiveness
  • Though deliberate inoculation of Artemia has been practised since the late 1970s (often in areas where no autochthonous Artemia was present), it is only recently that systematical studies are being undertaken on the spr...

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Pictures

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PictureTitleCaptionCopyright
Uterus of ovoviviparous Artemia filled with nauplii (first larvae are being released). (1) ovary with eggs.
TitleUterus
CaptionUterus of ovoviviparous Artemia filled with nauplii (first larvae are being released). (1) ovary with eggs.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Uterus of ovoviviparous Artemia filled with nauplii (first larvae are being released). (1) ovary with eggs.
UterusUterus of ovoviviparous Artemia filled with nauplii (first larvae are being released). (1) ovary with eggs.Laboratory of Aquaculture & Artemia Reference Center
Uterus of oviparous Artemia filled with cysts. (1) brown shell glands.
TitleUterus
CaptionUterus of oviparous Artemia filled with cysts. (1) brown shell glands.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Uterus of oviparous Artemia filled with cysts. (1) brown shell glands.
UterusUterus of oviparous Artemia filled with cysts. (1) brown shell glands.Laboratory of Aquaculture & Artemia Reference Center
Cyst in breaking stage; nauplius eye arrowed.
TitleCyst
CaptionCyst in breaking stage; nauplius eye arrowed.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Cyst in breaking stage; nauplius eye arrowed.
CystCyst in breaking stage; nauplius eye arrowed.Laboratory of Aquaculture & Artemia Reference Center
Embryo in "umbrella" stage (left) and instar I nauplius (right). (1) nauplius eye.
TitleEmbryo
CaptionEmbryo in "umbrella" stage (left) and instar I nauplius (right). (1) nauplius eye.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Embryo in "umbrella" stage (left) and instar I nauplius (right). (1) nauplius eye.
EmbryoEmbryo in "umbrella" stage (left) and instar I nauplius (right). (1) nauplius eye.Laboratory of Aquaculture & Artemia Reference Center
Instar V larva.  (1) nauplius eye; (2) lateral complex eye; (3) antenna; (4) labrum; (5) budding of thoracopods; (6) digestive tract.
TitleInstar V larva
CaptionInstar V larva. (1) nauplius eye; (2) lateral complex eye; (3) antenna; (4) labrum; (5) budding of thoracopods; (6) digestive tract.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Instar V larva.  (1) nauplius eye; (2) lateral complex eye; (3) antenna; (4) labrum; (5) budding of thoracopods; (6) digestive tract.
Instar V larvaInstar V larva. (1) nauplius eye; (2) lateral complex eye; (3) antenna; (4) labrum; (5) budding of thoracopods; (6) digestive tract.Laboratory of Aquaculture & Artemia Reference Center
Harvested Artemia, Vietnam.
TitleHarvested Artemia
CaptionHarvested Artemia, Vietnam.
CopyrightNguyen Van Hoa/Cantho University
Harvested Artemia, Vietnam.
Harvested ArtemiaHarvested Artemia, Vietnam.Nguyen Van Hoa/Cantho University
Head and thoracic region of young male: (1) antenna; (2) telopodite; (3) exopodite.
TitleHead and thoracic region
CaptionHead and thoracic region of young male: (1) antenna; (2) telopodite; (3) exopodite.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Head and thoracic region of young male: (1) antenna; (2) telopodite; (3) exopodite.
Head and thoracic regionHead and thoracic region of young male: (1) antenna; (2) telopodite; (3) exopodite.Laboratory of Aquaculture & Artemia Reference Center
Adult female.
TitleFemale Artemia
CaptionAdult female.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Adult female.
Female ArtemiaAdult female.Laboratory of Aquaculture & Artemia Reference Center
Head of an adult male. (1) antenna; (2) antennula; (3) lateral complex eye; (4) mandible.
TitleHead
CaptionHead of an adult male. (1) antenna; (2) antennula; (3) lateral complex eye; (4) mandible.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Head of an adult male. (1) antenna; (2) antennula; (3) lateral complex eye; (4) mandible.
HeadHead of an adult male. (1) antenna; (2) antennula; (3) lateral complex eye; (4) mandible.Laboratory of Aquaculture & Artemia Reference Center
Adult male.
TitleMale
CaptionAdult male.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Adult male.
MaleAdult male.Laboratory of Aquaculture & Artemia Reference Center
Detail of anterior thoracopods in adult Artemia. (1) exopodite; (2) telopodite; (3) endopodite.
TitleAnterior thoracopods
CaptionDetail of anterior thoracopods in adult Artemia. (1) exopodite; (2) telopodite; (3) endopodite.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Detail of anterior thoracopods in adult Artemia. (1) exopodite; (2) telopodite; (3) endopodite.
Anterior thoracopodsDetail of anterior thoracopods in adult Artemia. (1) exopodite; (2) telopodite; (3) endopodite.Laboratory of Aquaculture & Artemia Reference Center
Artemia couple in riding position. (1) uterus; (2) penis.
TitleArtemia couple
CaptionArtemia couple in riding position. (1) uterus; (2) penis.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Artemia couple in riding position. (1) uterus; (2) penis.
Artemia coupleArtemia couple in riding position. (1) uterus; (2) penis.Laboratory of Aquaculture & Artemia Reference Center
An artisanal salt/Artemia farm (Vietnam). Artemia biomass is collected in floating nets in the evaporation ponds.
TitleArtemia farm
CaptionAn artisanal salt/Artemia farm (Vietnam). Artemia biomass is collected in floating nets in the evaporation ponds.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
An artisanal salt/Artemia farm (Vietnam). Artemia biomass is collected in floating nets in the evaporation ponds.
Artemia farmAn artisanal salt/Artemia farm (Vietnam). Artemia biomass is collected in floating nets in the evaporation ponds.Laboratory of Aquaculture & Artemia Reference Center
Intensive Artemia culture.
TitleIntensive culture
CaptionIntensive Artemia culture.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Intensive Artemia culture.
Intensive cultureIntensive Artemia culture.Laboratory of Aquaculture & Artemia Reference Center
Energy content and dry weight of instar I, instar II, cold stored nauplii and decapsulated cysts (modified from Léger et al., 1987a).
TitleEnergy content and dry weight
CaptionEnergy content and dry weight of instar I, instar II, cold stored nauplii and decapsulated cysts (modified from Léger et al., 1987a).
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Energy content and dry weight of instar I, instar II, cold stored nauplii and decapsulated cysts (modified from Léger et al., 1987a).
Energy content and dry weightEnergy content and dry weight of instar I, instar II, cold stored nauplii and decapsulated cysts (modified from Léger et al., 1987a).Laboratory of Aquaculture & Artemia Reference Center
Typical setup for large scale hatching of Artemia.
TitleHatching
CaptionTypical setup for large scale hatching of Artemia.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Typical setup for large scale hatching of Artemia.
HatchingTypical setup for large scale hatching of Artemia.Laboratory of Aquaculture & Artemia Reference Center
Hatching tank after switching off aeration.
TitleHatching tank
CaptionHatching tank after switching off aeration.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Hatching tank after switching off aeration.
Hatching tankHatching tank after switching off aeration.Laboratory of Aquaculture & Artemia Reference Center
Concentrator/rinser used for an efficient harvest of large amounts of hatched Artemia.
TitleConcentrator/rinser
CaptionConcentrator/rinser used for an efficient harvest of large amounts of hatched Artemia.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Concentrator/rinser used for an efficient harvest of large amounts of hatched Artemia.
Concentrator/rinserConcentrator/rinser used for an efficient harvest of large amounts of hatched Artemia.Laboratory of Aquaculture & Artemia Reference Center
Principle of bio-encapsulation or enrichment.
TitleBio-encapsulation or enrichment
CaptionPrinciple of bio-encapsulation or enrichment.
CopyrightLaboratory of Aquaculture & Artemia Reference Center
Principle of bio-encapsulation or enrichment.
Bio-encapsulation or enrichmentPrinciple of bio-encapsulation or enrichment.Laboratory of Aquaculture & Artemia Reference Center

Identity

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Preferred Scientific Name

  • Artemia

Preferred Common Name

  • brine shrimp

Other Scientific Names

  • Artemia salina

International Common Names

  • Arabic: bahar-el-dud

Local Common Names

  • Germany: fezzan wurm
  • Spain: artemia
  • USA: sea monkeys

Summary of Invasiveness

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Though deliberate inoculation of Artemia has been practised since the late 1970s (often in areas where no autochthonous Artemia was present), it is only recently that systematical studies are being undertaken on the spread of Artemia beyond its original inoculation areas, and on the invasion of Artemia into suitable environments as a consequence of hatchery practices worldwide. Since both for hatchery and inoculation purposes A. franciscana is generally used, the invasion phenomenon in Artemia almost exclusively refers to spread of A. franciscana into an environment where this species did not occur naturally before, and where it enters into competition with an autochthonous population, if present. Though Artemia dispersal is generally linked to dispersion of cysts through wind and through migrating birds, not much has been documented so far on actual invasion patterns, but it can be assumed that these are linked to patterns of physiological tolerances and preferences of the competing populations. These patterns have been studied for areas where several species coexist naturally, and generally these studies refer to coexistence of a parthenogenetic and a bisexual strain.

In Artemia parthenogenetic strains (especially polyploid) often attain larger sizes and have faster growth rates than sexual diploid strains (Amat, 1980; Tobias et al., 1980). Evidence has been found that males subjected to a lower food regime have a lower survival rate than similarly reared females (Browne, 1980, 1982). Since fertilization is required for each new brood, high male mortality could lead to the domination of parthenogenetic populations in stressful habitats. In competition between sexual and parthenogenetic sympatric populations, the ability to respond to environmental stress may favour parthenogenesis (Browne and MacDonald, 1982). This factor, along with the theoretical two-fold increase in reproductive rate of parthenogenetic populations and the fact that only a single individual is required for colonisation (both of which may be important in many brine shrimp populations where temperature or desiccation cycles kill all but encysted individuals) may provide an explanation why parthenogenesis would be the predominant mode of reproduction in the Old World, or, at least, dominate over native Old World bisexual species (= non-franciscana) in the most stressful habitats.

Temporal cycling or niche partitioning would be expected based on temperature response profiles, as described by Browne et al. (1988). The effect of temperature on the performance of the population helps to determine the competitive interaction between bisexual and parthenogenetic brine shrimp strains and their relative success in competition experiments. Laboratory competition experiments result in the dominance of A. franciscana over parthenogenetic strains on one hand, and parthenogenetic populations over A. salina on the other hand (Browne, 1980; Browne and Halanych, 1989). Superior tolerance to high temperature of A. franciscana, as compared to various bisexual species, has been demonstrated (Vanhaecke and Sorgeloos, 1989). Nevertheless, different experimental conditions (in terms of temperature, salinity, food conditions) might lead to different results, and these laboratory results also need to be confirmed in the field. In most laboratory systems with other organisms competitive exclusion requires in the order of 10 to 100 generations, whereas for Artemia, exclusion occurs in only 2 or 3 generations (Lenz and Browne, 1991), which suggests a large or nearly complete niche overlap between the respective Artemia species.

For concrete examples of (natural) coexistence of different Artemia species on the Iberian peninsula, Iran and the People's Republic of China, see Van Stappen (2002).

During the 1980s, Artemia were intentionally introduced into several (sub)-tropical countries including Thailand, Philippines, Vietnam, Indonesia, China, India, Iran, Egypt, Sri Lanka and Panama from the USA (Vanhaecke et al., 1987; Lavens and Sorgeloos, 1996). However, Artemia has also spread unintentionally beyond the original inoculation areas and/or has invaded new territories in France, Portugal, Spain, China, India, Egypt and the United Arab Emirates (Narciso, 1989; Thiery and Robert, 1992; Amat, 2002; Van Stappen, 2002; Amat et al., 2003a, b; Xin Naihong, 2003)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Crustacea
  •                 Class: Branchiopoda
  •                     Order: Anostraca
  •                         Family: Artemiidae
  •                             Genus: Artemia

Description

Top of page Biology and Ecology

Artemia are found in isolated habitats, namely natural salt lakes and man-made salterns, in temperate to tropical regions. The distribution of these sites over the continents is very uneven, mainly reflecting sampling and exploration activities (see Pictures). The occurrence of Artemia is constrained, on one hand, to sites where conditions allow the animals to survive throughout the year, and on the other hand, to regions where the seasonality of the environment is stable (Lenz, 1987; Amat et al., 1995).

The common feature of all Artemia biotopes is high salinity. Brine shrimps do not possess any anatomical defense mechanism against predation but they have developed unique physiological adaptations to high salinity which provides a very efficient ecological defense against predation as they can thrive in salinities that are lethal to any common aquatic predator. These adaptations consist of:

  • a very efficient osmoregulatory system
  • the capacity to synthesize very efficient respiratory pigments to cope with the low O2 levels at high salinities
  • the ability to produce dormant cysts when environmental conditions endanger the survival of the species.

Artemia is incapable of active dispersion, therefore wind and waterfowl (especially flamingos) are the most important natural dispersion vectors (apart from inoculation by man). The floating cysts adhere to feet and feathers of birds and, when ingested, they remain intact for at least a couple of days in the digestive tract of birds.

Life Cycle

Under favourable conditions, fertilized eggs develop into free-swimming nauplii (= ovoviviparous reproduction) (see Pictures), which are released by the mother. In extreme conditions (e.g. too high or too low salinity, low oxygen levels) the embryos only develop up to the gastrula stage. At this point they are surrounded by a thick shell, enter a state of metabolic dormancy (diapause) and are then released by the female (= oviparous reproduction) (see Pictures).

In principle, both oviparity and ovoviviparity are found in all Artemia strains, and females can switch reproductive modes from one ovulation to the next. The cysts usually float in the high salinity waters and are blown ashore where they accumulate and dry. As a result of this dehydration process the diapause mechanism is generally inactivated; cysts are now in a state of quiescence and can resume further embryonic development when hydrated in optimal hatching conditions. Under optimal conditions brine shrimp can live for several months, grow from nauplius to adult in only 8 days and reproduce at a rate of up to 300 nauplii or cysts every 4 days.

Strain-specific Characteristics

In contrast to nutritional quality, cyst diameter and resistance to high temperature are considered strain-specific and remain relatively constant (Vanhaecke and Sorgeloos, 1980), i.e. they have become genotypical as a result of long-term adaptations of the strain to the local conditions.

In general cyst diameter of the same strain remains constant in spite of small variations between batches, possibly caused by environmental and/or processing factors. Other biometrical characteristics such as cyst volume, cyst dry weight, instar 1 naupliar length, individual naupliar weight and naupliar volume, energy content etc., show a high correlation with the cyst diameter (Vanhaecke et al., 1983). As a consequence, biometrical parameters, in particular cyst diameter, are good tools to characterize Artemia strains, and to help to define the origin of unknown or even mixed cyst samples (Vanhaecke and Sorgeloos, 1980). Some general correlations can also be made between sibling species and size:

  • parthenogenetic Artemia produce large cysts
  • A. salina
  • produce large cysts with a thick chorion
  • A. franciscana
  • and A. persimilis produce small or intermediate cysts with a thin chorion.

Comparative studies of hatching behaviour of cysts of different origin show a considerable variation in hatching percentage, rate and efficiency (Vanhaecke and Sorgeloos, 1982; 1983). However, none of these parameters is strain-specific as they are influenced by a wide array of factors such as harvesting, processing, storage and hatching techniques, as well as production conditions affecting the parental generation.

Both temperature and salinity affect survival and growth, the effect of temperature being more pronounced (Vanhaecke et al., 1984). For most commonly used Artemia the optimal temperature range is 20-25°C. There seems to be some genetic adaptation to high temperatures (Clegg et al., 2001).

Life history and reproductive characteristics of Artemia strains are important factors when an introduction of brine shrimp to a new habitat is considered, especially when competition with a local strain is to be expected. These competitive abilities are related to factors such as the length of reproductive, pre- and post-reproductive periods, total lifespan, number of offspring per brood, broods per female, interval between broods etc. In general, New World (zygogenetic) populations have a very large number of offspring per brood, a large number of offspring/day/female and a fast development time to sexual maturity. These are all favourable characteristics when compared with those of Old World zygogenetic and parthenogenetic Artemia (Browne et al., 1984; 1991).

Age at first reproduction is a key factor determining the population growth rate, and the rate of colonisation of new environments with limited nutrient resources. Consequently, if environmental preferences and nutritional factors do not interfere, New World zygogenetic species generally outcompete parthenogenetic strains, the latter in their turn predominating over Old World zygogenetic species. Inoculation experiments in natural habitats therefore require prior screening of candidate strains and of indigenous local populations, as well as the study of prevailing environmental conditions.

Morphology

Under certain conditions, Artemia produces cysts that float on the water surface and that are driven ashore by wind and waves. These cysts are metabolically inactive and do not develop further as long as they are kept dry. Upon immersion in seawater, the biconcave-shaped cysts hydrate, become spherical and within the shell, the embryo resumes its interrupted metabolism. After about 20 h the outer membrane of the cyst bursts (= "breaking") and the embryo appears, surrounded by the hatching membrane (see Pictures). While the embryo hangs underneath the empty shell (= "umbrella" stage) the development of the nauplius is completed and within a short period of time the hatching membrane is ruptured (= "hatching") giving rise to the free-swimming nauplius (see Pictures).

The first larval stage (instar 1, 400 to 500 µm in length) has a brownish-orange colour, a red nauplius eye in the head region and three pairs of appendages, i.e. the first antennae (sensorial function), the second antennae (locomotory and filter-feeding function) and the mandibles (food uptake function). The ventral side is covered by a large labrum (food uptake and transfer of particles from the filtering setae into the mouth). The instar 1 larva does not take up food as its digestive system is not yet functional; it relies completely on its yolk reserves. After about 8 h, the animal moults into the second larval stage (instar 2). Small food particles (e.g. algal cells, bacteria, detritus) ranging in size from 1 to 50 µm are filtered out by the second antennae and ingested into the now functional digestive tract. The larva grows and differentiates through a number of moults: one naupliar, four metanauplia, seven postmetanauplia and five postlarval stages have been described (Hentschel, 1968; Schrehardt, 1987).

Paired lobular appendages appear in the trunk region and differentiate into thoracopods (see Pictures). On both sides of the nauplius eye lateral complex eyes begin to develop (see Pictures). From the tenth instar stage on, important morphological as well as functional changes begin to take place i.e. the antennae lose their locomotory function and undergo sexual differentiation. In males (see Pictures) they develop into hooked graspers, whereas the female antennae degenerate into sensorial appendages (see Pictures). The thoracopods are now differentiated into three functional parts (see Pictures): i.e. the telopodites and endopodites (locomotory and filter feeding), and the membranous exopodites (gills).

Adult Artemia are typical primitive arthropods (8-12 mm in length) having an elongated segmented body with two stalked complex eyes, a linear digestive tract, sensorial antennulae, a pair of functional thoracopods on each of the 11 thoracic segments, and a furca on the last abdominal segment. The entire body is covered with a thin, flexible exoskeleton of chitin to which muscles are attached internally. The male has a paired penis on the first of the eight abdominal segments (see Pictures). Female Artemia can easily be recognized by the brood pouch or uterus situated in the same segment, just behind the eleventh pair of thoracopods (see Pictures). The female reproductive system consists of ovaries and oviducts leading into the single, median uterus wherein several clusters of shell glands open. The ovaries are paired tubular structures extending into the abdomen (see Pictures). Adult females ovulate approximately every 140 hours, depending on rearing conditions and whether development of embryos occurs oviparously or ovoviviparously. In females, spawning is followed by a moult, after which ovulation takes place. The oviducts emerge from the ovaries near the anterior part of the third abdominal segment (Cassel, 1937). Each oviduct empties into the antero-lateral border of the uterus. The lateral pouches function as seminal receptacles during the time between copulation and fertilization (within 1 hour) (Criel, 1980ab; Benesch, 1969). Once ripe, the eggs developing in the ovaries become spherical and migrate via two oviducts into the unpaired uterus.

Distribution

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Artemia populations are found in about 500 natural salt lakes and man-made salterns scattered throughout the tropical, subtropical and temperate climatic zones, along coastlines as well as inland (see Pictures). The distribution of these sites over the continents is very uneven, mainly reflecting sampling and exploration activities. As such it does not give a precise picture of the actual global occurrence of Artemia. A continued survey will undoubtedly lead to the discovery of many more Artemia biotopes in different parts of the world. Lists of Artemia sites principally include biotopes where the population reappears each year. Additionally, temporal populations occur where Artemia is introduced on a seasonal basis, mostly in seasonal salt operations, e.g. Panama, Costa Rica, Myanmar, Thailand, Philippines, Vietnam, Indonesia and other places (Vanhaecke et al., 1987). Eventually, depending on the climatic conditions and management procedures, some of these populations may become established as feral strains; e.g. Cam Ranh Bay, Vietnam (Vu Do Quynh and Nguyen Ngoc Lam, 1987). Artemia is also present in Kyrgyzstan and Tajikstan (L Nagorskaya, Institute of Zoology, Belarus, personal communication 2004).

Two critical factors determine the population dynamics of Artemia and its biogeographical distribution: firstly, whether water body conditions allow the animals to survive throughout the year, and secondly, whether the seasonality of the environment is predictable or not (Lenz, 1987; Amat et al., 1995). The common feature of all Artemia biotopes is their high salinity. Salinity is without doubt the predominant abiotic factor determining the presence of Artemia and consequently limiting its geographical distribution. Other variables (temperature, light intensity and primary food production) may have an influence on the quantitative aspects of the Artemia population, or may cause only a temporary absence of brine shrimp.

The brine shrimp Artemia comprises a group of zygogenetic and parthenogenetic, morphologically similar, populations. Speciation in the genus should be regarded as a complex, multi-dimensional process involving a variety of environmental and genomic factors. The identification of zygogenetic Artemia species has been established by a multidisciplinary approach, including cross-breeding tests, morphological differentiation, cytogenetics, allozyme studies, nuclear and mitochondrial DNA sequencing. With the exception of cross-mating, all these techniques have also contributed to identifying the parthenogenetic types described as A. parthenogenetica by Barigozzi (1974) as well as to gain insights on population structure, origin and amount of clonal diversity.

Endemic to the Old World are the parthenogenetic types designated by Barigozzi (1974) as A. parthenogenetica (with different levels of ploidy, found in Europe, Africa, Asia and Australia), the zygogenetic A. salina Leach 1819 (Mediterranean area) (Triantaphyllidis et al., 1997), A. urmiana (Günther, 1890) (Iran), A. sinica (Cai, 1989) (continental China), Artemia sp. (Pilla and Beardmore, 1994) (Kazakhstan), and A. tibetiana (Abatzopoulos et al., 1998, 2002) (Tibet). Endemic to the New World are A. persimilis (Piccinelli and Prosdocimi, 1968) (Argentina) and A. franciscana (Kellogg, 1906) (North, Central and South America), with A. (franciscana) monica being a special case of a population described for an ecologically unique habitat (Mono Lake, USA).


Suitable Habitat Conditions

As a result of the introduction of A. franciscana in solar saltworks for improved salt production and/or for harvesting cysts and biomass for use in the aquaculture industry, permanent populations of this species are nowadays found in Brazil, Australia, China, Egypt, Portugal etc. Seasonal A. franciscana farming is practised in many tropical and subtropical countries such as the Philippines, Thailand, Vietnam, Sri Lanka (Triantaphyllidis et al., 1998). However, due to the particular climatic conditions of these countries, the Artemia populations are not permanent and annual inoculations are required.

Two critical factors determine the population dynamics of Artemia and its zoogeographical distribution: firstly, whether water body conditions allow the animals to survive throughout the year, and secondly, whether the seasonality of the environment is predictable or not. A few general remarks can be made regarding the impact of some environmental factors on the Artemia distribution.


Salinity


Salinity is without any doubt the predominant abiotic factor determining the presence of Artemia and consequently limiting its geographical distribution. Salinity as a crucial factor for Artemia presence is linked to the occurrence of predators of brine shrimp at lower and intermediate salinities. Other variables (temperature, light intensity and primary food production) may have an influence on the quantitative aspects of the Artemia population, or may cause only a temporary absence of brine shrimp. On the other hand, not all highly saline biotopes are populated with Artemia. Vanhaecke et al. (1987) have used the classification system of Thornthwaite (1948) to relate the natural distribution of Artemia sites to different climate types. This study revealed that the geographical distribution of brine shrimp is limited by climatological conditions, i.e. no natural Artemia populations are found in humid climate types, and 97% of the biotopes are located in areas where yearly evaporation exceeds yearly precipitation. In humid climate types, normally human intervention is required to keep high salinity levels (e.g. removal of freshwater stratification layers by drainage or pumping, storage of brine in reservoirs). Additionally, Vanhaecke et al. (1987) outlined the potential distribution pattern of Artemia on a global level, stressing nevertheless the restricted relevance of this kind of extrapolation: climate types may vary significantly even within relatively short distances and specific local conditions may result in isolated microclimates suitable or unsuitable for Artemia. Moreover, besides general annual precipitation and evaporation figures, seasonal distribution of precipitation and evaporation are also important, and more detailed climatological conditions and water balance data have to be used to obtain an accurate idea about the potential world distribution of Artemia.

Brine shrimp have been found alive in supersaturated brines at salinities as high as 340 ppt (Post and Youssef, 1977). Under these extreme conditions, however, the animals barely manage to survive and their normal physiological and metabolic functions are seriously affected. The lower salinity limit in nature is basically a function of the presence of predatory animals. Brine shrimp are rarely found in waters with salinity lower than 45 ppt, although physiologically they thrive in seawater and even in brackish waters (Persoone and Sorgeloos, 1980). Brine shrimp do not possess any anatomical defense mechanism against predation and consequently Artemia populations are always in danger at salinities tolerated by carnivorous species. The list of Artemia predators includes by definition all species populating natural seawaters and feeding on zooplankton. As a general rule, the lowest salinity at which Artemia is found in nature thus varies from place to place and is determined by the upper salinity tolerance level of the local predator(s).

No clear optimum can be defined for salinity of the Artemia environment. For physiological reasons this optimum must be situated towards the lower end of the salinity range, as higher ambient salinity requires higher energy costs for osmoregulation. Ambient salinity finally also plays a role in cyst metabolism, as Artemia cysts will only start to develop when the salinity of the medium drops below a certain threshold value, which is strain dependent. At salinities above this threshold Artemia cysts will never hatch because the hydration level they reach is insufficient, which is one of the prerequisites for the onset of the hatching metabolism (Lavens and Sorgeloos, 1987). Situations of a temporary low salinity often occur in a salt lake, e.g. in a restricted area of inflowing freshwater, or after rainfall when for a while a freshwater layer remains on top of the heavier salt water.


Ionic Composition


Artemia can withstand environments in which the ratio of the major anions and cations may be totally different from that in seawater, and even reach extremely high or low values in comparison to natural seawater (Persoone and Sorgeloos, 1980). Depending on the prevailing anions, Artemia may thus inhabit chloride, sulphate or carbonate waters and/or combinations of two or even three major anions.


Temperature


Apart from salinity, temperature also affects the distribution pattern of Artemia (Vanhaecke et al., 1987). No Artemia is found in the cold tundra or frost climate areas, as the year-round prevailing extremely low temperatures preclude Artemia development. Moreover, potential evaporation is very limited in these regions, which (with only very few exceptions) excludes the presence of highly saline biotopes. A lot of strains are found in continental areas (continental USA and China, Central Asia, South Siberia) with extremely cold winter temperatures, but hot summer temperatures allow the hatching of the cysts and subsequent colonisation of the environment.

The maximum temperature that Artemia populations tolerate has repeatedly been reported to be close to 35°C, a temperature often attained in the shallow tropical salterns that constitute a large part of the Artemia habitats. This tolerance threshold is, however, strain-dependent. Physiological adaptation of Artemia after a number of generations to the high temperatures (± 40°C) in Vietnamese salt ponds has been reported (Clegg et al., 2001). As for salinity, temperature optima are difficult to define and are strain-dependent; generally, however, one can state that the optimum for Artemia biomass must be situated in the range 25-30°C.


Biotic Elements


While hypersaline environments are characterised by monocultures of Artemia as major zooplankton, lower and intermediate salinity habitats are populated by various groups of invertebrates. Depending on the seasonal cycle or the salinity gradient in certain biotopes (e.g. in man-managed saltworks) there can thus be a range where there is some degree of coexistence with brine shrimp. Numerous studies report about the faunal and floral elements of these biotopes, and the extent to which they are found in coexistence with Artemia. For more information about this and other topics related to Artemia zoogeography, see the review by Van Stappen (2002).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ChinaPresentPresent based on regional distribution.
-FujianPresentNativeVan Stappen, 2002
-GansuPresentNativeVan Stappen, 2002
-GuangdongPresentNativeVan Stappen, 2002
-HainanPresentNativeVan Stappen, 2002
-HubeiPresentNativeVan Stappen, 2002
-JiangsuPresentNativeVan Stappen, 2002
-LiaoningPresentNativeVan Stappen, 2002
-Nei MengguPresentNativeVan Stappen, 2002
-NingxiaPresentNativeVan Stappen, 2002
-QinghaiPresentNativeVan Stappen, 2002
-ShaanxiPresentNativeVan Stappen, 2002
-ShandongPresentNativeVan Stappen, 2002
-ShanxiPresentNativeVan Stappen, 2002
-TibetPresentNativeVan Stappen, 2002
-XinjiangPresentNativeVan Stappen, 2002
-ZhejiangPresentNativeVan Stappen, 2002
IndiaPresentNative Invasive Van Stappen, 2002
IndonesiaPresentIntroducedLavens and Sorgeloos, 2000
IranPresentNativeVan Stappen, 2002
IraqPresentNativeVan Stappen, 2002
IsraelPresentNativeVan Stappen, 2002
JapanPresentNativeVan Stappen, 2002
KazakhstanPresentNativeVan Stappen, 2002
Korea, Republic ofPresentNativeVan Stappen, 2002
KuwaitPresentNativeVan Stappen, 2002
MalaysiaPresentIntroducedLavens and Sorgeloos, 2000
MongoliaPresentNativeVan Stappen, 2002
MyanmarPresentIntroducedVanhaecke et al., 1987
PakistanPresentNativeVan Stappen, 2002
PhilippinesPresentIntroducedLavens and Sorgeloos, 2000
Saudi ArabiaPresentNativeVan Stappen, 2002
Sri LankaPresentNativeVan Stappen, 2002
SyriaPresentNativeVan Stappen, 2002
TaiwanPresentNativeVan Stappen, 2002
ThailandPresentIntroducedLavens and Sorgeloos, 2000
TurkeyPresentNativeVan Stappen, 2002
TurkmenistanPresentNativeVan Stappen, 2002
United Arab EmiratesPresentNativeVan Stappen, 2002
UzbekistanPresentNativeVan Stappen, 2002
VietnamPresentIntroducedLavens and Sorgeloos, 2000

Africa

AlgeriaPresentNativeVan Stappen, 2002
Cape VerdePresentNativeVan Stappen, 2002
EgyptPresentNativeVan Stappen, 2002
EritreaPresentNativeLavens and Sorgeloos, 2000
KenyaPresentNativeVan Stappen, 2002
LibyaPresentNativeVan Stappen, 2002
MadagascarPresentNativeVan Stappen, 2002
MoroccoPresentNativeAmat, 2002; Van Stappen, 2002
MozambiquePresentNativeVan Stappen, 2002
NamibiaPresentNativeVan Stappen, 2002
NigerPresentNativeVan Stappen, 2002
SenegalPresentNativeVan Stappen, 2002
South AfricaPresentNativeVan Stappen, 2002
Spain
-Canary IslandsPresentNativeVan Stappen, 2002
TunisiaPresentNativeVan Stappen, 2002

North America

CanadaPresentPresent based on regional distribution.
-AlbertaPresentNativeVan Stappen, 2002
-SaskatchewanPresentNativeVan Stappen, 2002
MexicoPresentNativeVan Stappen, 2002
USAPresentPresent based on regional distribution.
-ArizonaPresentNativeVan Stappen, 2002
-CaliforniaPresentNativeVan Stappen, 2002
-HawaiiPresentNativeVan Stappen, 2002
-NebraskaPresentNativeVan Stappen, 2002
-NevadaPresentNativeVan Stappen, 2002
-New MexicoPresentNativeVan Stappen, 2002
-North DakotaPresentNativeVan Stappen, 2002
-OregonPresentNativeVan Stappen, 2002
-TexasPresentNativeVan Stappen, 2002
-UtahPresentNativeVan Stappen, 2002
-WashingtonPresentNativeVan Stappen, 2002

Central America and Caribbean

Antigua and BarbudaPresentNativeVan Stappen, 2002
ArubaPresentNativeVan Stappen, 2002
BahamasPresentNativeVan Stappen, 2002
British Virgin IslandsPresentNativeVan Stappen, 2002
Costa RicaPresentNativeVan Stappen, 2002
CuraçaoPresentNativeVan Stappen, 2002
Dominican RepublicPresentNativeVan Stappen, 2002
HaitiPresentNativeVan Stappen, 2002
JamaicaPresentNativeVan Stappen, 2002
Netherlands AntillesPresentNativeVan Stappen, 2002
NicaraguaPresentNativeVan Stappen, 2002
PanamaPresentIntroducedVanhaecke et al., 1987
Puerto RicoPresentNativeVan Stappen, 2002
Saint Kitts and NevisPresentNativeVan Stappen, 2002
Turks and Caicos IslandsPresentNativeVan Stappen, 2002

South America

ArgentinaPresentNativeVan Stappen, 2002
BoliviaPresentNativeVan Stappen, 2002
BrazilPresentPresent based on regional distribution.
-CearaPresentNativeVan Stappen, 2002
-Rio Grande do NortePresentNativeVan Stappen, 2002
ChilePresentNativeVan Stappen, 2002
ColombiaPresentNativeVan Stappen, 2002
EcuadorPresentNativeVan Stappen, 2002
PeruPresentNativeVan Stappen, 2002
VenezuelaPresentNativeVan Stappen, 2002

Europe

BulgariaPresentNativeVan Stappen, 2002
CroatiaPresentNativeVan Stappen, 2002
CyprusPresentNativeVan Stappen, 2002
FrancePresentNativeVan Stappen, 2002
GreecePresentNativeVan Stappen, 2002
ItalyPresentNativeVan Stappen, 2002
PortugalPresentNativeVan Stappen, 2002
RomaniaPresentNativeVan Stappen, 2002
Russian FederationPresentPresent based on regional distribution.
-Eastern SiberiaPresentNativeVan Stappen, 2002
-Southern RussiaPresentNativeVan Stappen, 2002
-Western SiberiaPresentNativeVan Stappen, 2002
SpainPresentNativeVan Stappen, 2002
-Balearic IslandsPresentNativeVan Stappen, 2002
UkrainePresentNativeVan Stappen, 2002
Yugoslavia (Serbia and Montenegro)PresentNativeVan Stappen, 2002

Oceania

AustraliaPresentPresent based on regional distribution.
-QueenslandPresentNativeVan Stappen, 2002
-South AustraliaPresentNativeVan Stappen, 2002
-Western AustraliaPresentNativeVan Stappen, 2002
New ZealandPresentIntroducedVan Stappen, 2002

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
China USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
Egypt USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
India USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisationLavens and Sogeloos (1996); Vanhaecke et al. (1987)
Indonesia USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
Iran USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
Panama USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
Philippines USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
Sri Lanka USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
Thailand USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)
Vietnam USA 1980s Aquaculture (pathway cause)Government|Private sector|International organisation Yes Yes Lavens and Sogeloos (1996); Vanhaecke et al. (1987)

Climate

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ClimateStatusDescriptionRemark
A - Tropical/Megathermal climate Tolerated Average temp. of coolest month > 18°C, > 1500mm precipitation annually
B - Dry (arid and semi-arid) Preferred < 860mm precipitation annually

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 18 35
Mean maximum temperature of hottest month (ºC) 35
Mean minimum temperature of coldest month (ºC) 18

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Ammonia [unionised] (mg/l) >50 Harmful Adult
Dissolved oxygen (mg/l) <2 Harmful Adult
Dissolved oxygen (mg/l) 5 Optimum Adult
Illumination (Lux illuminance) >0 Harmful Egg
Illumination (Lux illuminance) 0 Optimum Egg
Illumination (Lux illuminance) 32 Harmful Adult
Nitrate (mg/l) >1000 Harmful Adult
Nitrite (mg/l) >320 Harmful Adult
Salinity (part per thousand) 30 Optimum Adult strain specific
Salinity (part per thousand) <25 >250 Harmful Adult strain specific
Water pH (pH) 8 Optimum Adult strain specific
Water pH (pH) <6.5 >8 Harmful Adult strain specific
Water temperature (ºC temperature) -18 Optimum Egg strain specific, optimum spawning temperature 25°C
Water temperature (ºC temperature) 25 Optimum Adult strain specific, optimum spawning temperature 25°C
Water temperature (ºC temperature) <-18 >40 Harmful Egg strain specific, optimum spawning temperature 25°C
Water temperature (ºC temperature) <15 >32 Harmful Adult strain specific, optimum spawning temperature 25°C

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aphanius Adult/Larval
Ardeidae All Stages
Corixidae Adult/Larval
Phoenicopteridae All Stages
Recurvirostra All Stages

Environmental Impact

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An inoculation in 1977 in the Macau saltworks, Natal, northeastern Brazil, with Artemia from San Francisco Bay, USA, was extremely successful, and brine shrimp soon spread out over 3000 ha of saltworks (Persoone and Sorgeloos, 1980). As a result of the introduction of A. franciscana in solar saltworks for improved salt production and/or for harvesting cysts and biomass for use in the aquaculture industry, permanent and temporary (to be re-inoculated each dry season) populations of this species are nowadays found worldwide.

It is widely recognized that salt lakes are unique and well-balanced ecosystems, and that there are differences in their fauna between continents and regions, and even locally, according to their salinity fluctuations, water permanence, seasonality etc. Many salt lakes are of intermediate or small size, and this very specific environment, including the food chain with Artemia and waterfowl is very vulnerable for deterioration by human intervention. Threats to salt lakes are numerous, and mainly consist of desiccation by drainage or diversion of influents, or pollution. For coastal saltworks, urbanization projects (industry, harbour infrastructure, tourism, expansion of residential areas) represent a supplementary threat, especially in industrialized or industrializing countries.

Although the practice of Artemia introduction frequently ensures social and economic benefits, particularly in developing countries, it also bears certain risks (Beardmore, 1987; Beardmore et al., 1997). An obvious effect is that competition with local (or near-by) strains or species of Artemia may occur (Geddes and Williams, 1987), which may lead to the extinction of some genotypes, or at worst, of one of the competitors. Competition experiments suggest that A. franciscana may outcompete others (Browne, 1980; Browne and Halanych, 1989). The effect of one introduction will not remain local but may have consequences over large areas (such as Natal). Amat et al. (1995) and Narciso (1989) report about the presence of A. franciscana populations in the Iberian Peninsula, where this allochthonous strain, due to intentional or non-intentional inoculations (e.g. through hatchery effluents), in some locations has outcompeted the local Artemia populations.

Crosses have often been performed in laboratory-reared populations, e.g. of A. franciscana and A. persimilis (Gajardo et al., 2001). The production of laboratory hybrids between morphologically or genetically divergent allopatric populations appears to be a common phenomenon in some Artemia populations (Bowen et al., 1985; Pilla and Beardmore, 1994), e.g. between eastern Old World sexual populations. This might be related to the fact that these populations have only completed the first stage of the allopatric speciation process (i.e. geographical separation) but have not completed the second stage required for the development of pre-mating isolating mechanisms (for more genetic background on cross-fertility vs. reproductive isolation of different Artemia species, see Gajardo et al., 2002). Nevertheless, hybridization between different species in nature, due to deliberate or undeliberate introduction by man, has not been documented so far: the species introduced, generally A. franciscana, is genetically too distant from the local sexual populations (e.g. A. salina), or the local populations are often parthenogenetic.

Measures to ensure that the Artemia biodiversity is conserved include the establishment of gene banks (cysts), close monitoring of inoculation policies, and where possible the use of indigenous Artemia for inoculating Artemia-free waters (Beardmore, 1987).

Another threat for Artemia populations worldwide is posed by the massive harvesting practices, that have been going on for a few decades (e.g. in the Great Salt Lake, USA) or that have been launched in recent years at several sites at an intensified rate in view of a menacing cyst shortage. It is largely speculative whether these indiscriminate harvests actually endanger the habitat’s standing crop; Artemia habitats differ enormously in size and in productivity. However, some degree of selection on the local gene pool (e.g. favouring ovoviviparous reproduction) may be imposed by systematic long-term harvesting of Artemia cysts.

Uses List

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Animal feed, fodder, forage

  • Live feed

References

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Abatzopoulos T; Zhang B; Sorgeloos P, 1998. Artemia tibetiana: preliminary characterization of a new Artemia species found in Tibet (People’s Republic of China) International Study on Artemia LIX. Int. J. Salt Lake Res, 7:1-44.

Abelin P; Tackaert W; Sorgeloos P, 1991. Ensiled Artemia biomass: a promising and practical feed for penaeid shrimp postlarvae. In: Lavens P, Sorgeloos P, Jaspers E, Ollevier F, eds. Larvi’91 - Fish & Crustacean Larviculture Symposium. Ostend, Belgium: European Aquaculture Society, Special Publication No. 15, 125-127.

Amat F, 1980. Differentiation in Artemia strains from Spain. In: Persoone G, Sorgeloos P, Roels O, Jaspers E, eds. The brine shrimp Artemia, Vol. 2. Belgium: Universa Press, 19-39.

Amat F, 2002. Artemia biodiversity: current global resources and their sustainable exploitation. INCO Workshop on Artemia Biodiversity, Beijing, China, September 23-26, 2002.

Amat F; Barata C; Hontoria F; Navarro JC; Varó I, 1995. Biogeography of the genus Artemia (Crustacea, Branchiopoda, Anostraca) in Spain. Int. J. Salt Lake Res, 3:175-190.

Amat F; Baxevanis AD; Triantafyllidis A; Tzika A; Mura G; Abatzopoulos TJ, 2003. The use of RFLP 16S rDNA analysis for detecting invasion of Artemia franciscana in Western Mediterranean region. INCO Workshop on Artemia Biodiversity, Puerto Varas, Chile, November 17-19, 2003.

Amat F; Hontoria F; Ruiz O; Sánchez M; Green A; Figuerola J; Hortas F, 2003. Artemia franciscana como especie exótica invasora en el Mediterráneo Occidental. I Congreso Nacional sobre Especies Exóticas Invasoras. (EEI 2003). Universidad de Leon, Spain, 4 - 7 June 2003.

Anbaya Almalul MA, 2000. Effect of temperature and density on storage characteristics of Artemia nauplii. MSc thesis. Ghent, Belguim: Faculty for Agricultural and Applied Biological Sciences, Ghent University , 1-52.

Baert P; Anh NTN; Vu Do Quynh; Hoa NV; Sorgeloos P, 1997. Increasing cyst yields in Artemia culture ponds in Vietnam: the multi-cycle system. Aquaculture Research, 28(10):809-814.

Barigozzi C, 1974. Artemia: A survey of its significance in genetic problems. Evolutionary Biology, 7:221-251.

Beardmore JA, 1987. Concluding remarks for Symposium Session I: Morphology, Ecotoxicology, Radiobiology, Genetics. In: Sorgeloos P, Bengtson DA, Decleir W, Jaspers E, eds. Artemia Research and its Applications, Vol. 1. Wetteren, Belgium: Universa Press, 345-346.

Beardmore JA; Mair GC; Lewis RI, 1997. Biodiversity in aquatic systems in relation to aquaculture. Aquaculture Research, 28(10):829-839.

Belk D; Brtek J, 1995. Checklist of the Anostraca. Hydrobiologia, 298(1/3):315-353.

Benesch R, 1969. Zur Ontogenie und Morphologie von Artemia salina L., Zool. Jb. Anat. Bd, 86:307-458.

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Bowen ST; Fogarino EA; Hitchner KN; Dana GL; Chow VHS; Buoncristiani MR; Carl JR, 1985. Ecological isolation in Artemia: population differences in tolerance of anion concentrations. Journal of Crustacean Biology, 5:106-129.

Brown RA; Bowen ST, 1991. Taxonomy and population genetics of Artemia.. Artemia biology., 221-235.

Browne RA, 1980. Competition experiments between parthenogenetic and sexual strains of the brine shrimp, Artemia salina. Ecology, 61(3):471-474.

Browne RA; Davis LE; Sallee SE, 1988. Effects of temperature and relative fitness of sexual and asexual brine shrimp Artemia. Journal of Experimental Marine Biology and Ecology, 124:1-20.

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Browne RA; MacDonald GH, 1982. Biogeography of the brine shrimp, Artemia: distribution of parthenogenetic and sexual populations. Journal of Biogeography, 9:331-338.

Browne RA; Sallee SE, 1984. Partitioning genetic and environmental components of reproduction and lifespan in Artemia.. Ecology, 65(3):949-960.

Cai Y, 1989. A redescription of the brine shrimp (Artemia sinica). Wasman J. Biol, 7:221-251.

Cassel JD, 1937. The morphology of Artemia salina (Linnaeus). MA Thesis. California USA: Leland Stanford Junior University, 108.

Chair M; Nelis HJ; Léger P; Sorgeloos P; De Leenheer A, 1996. Accumulation of trimethoprim, sulfamethoxazole, and N-acetylsulfamethoxazole in fish and shrimp fed medicated Artemia franciscana. Antimicrobial Agents and Chemotherapy, 40:1649-1652.

Chen JC; Chen KJ; Liao JM, 1989. Joint action of ammonia and nitrite on Artemia nauplii. Aquaculture, 77:329-336.

Clegg JS; Nguyen Van Hoa and Sorgeloos P, 2001. Thermal tolerance and heat shock proteins in encysted embryos of Artemia from widely different thermal habitats. Hydrobiologia, 466:221-229.

Correa Sandoval F; Cordero Esquivel B; Valenzuela-Espinoza E; Escobar Fernandez R, 1994. Biochemical composition of laboratory-cultured adults of Artemia franciscana Kellogg 1906. Rivista Italiana di Aquacoltura 29:63-66.

Correa Sandoval F; Ramirez LFB; Lobina DV, 1993. The biochemical composition of the cysts of some Mexican populations of Artemia franciscana Kellogg 1906. Comparative Biochemistry and Physiology B 104:163-167.

Coutteau P; Sorgeloos P, 1989. Feeding of the brine shrimp Artemia on yeast: effect of mechanical disturbance, animal density, water quality and light intensity. In: Aquaculture Europe 1989 - Book of Abstracts. Bredene, Belgium: EAS Special Publication No.10:75-76.

Criel G, 1980. Morphology of the genital apparatus of Artemia: a review. In: Persoone G, Sorgeloos P, Roels O, Jaspers E, eds. The brine shrimp Artemia, Vol. I, Morphology, Genetics, Radiobiology, Toxicology. Wetteren, Belgium: Universa Press, 75-86.

Criel G, 1980. Ultrastructural observations on the oviduct of Artemia. In: Persoone G, Sorgeloos P, Roels O, Jaspers E, eds. The brine shrimp Artemia, Vol. I, Morphology, Genetics, Radiobiology, Toxicology. Wetteren, Belgium: Universa Press, 87-95.

D’Agostino A, 1980. The vital requirements of Artemia: physiology and nutrition. In: Persoone G, Sorgeloos P, Roels O, Jaspers E, eds. The brine shrimp Artemia. Vol 2.: Physiology, Biochemistry, Molecular Biology. Wetteren, Belgium Universa Press, 55-82.

Dehasque M; Devresse B; Sorgeloos P, 1993. Effective suppression of bacterial bloom during hatching and enrichment of Artemia and its applicability in fish/shrimp hatcheries. In: Book of Abstracts of tha World Aquaculture ‘93. Ostend, Belgium: European Aquaculture Society, Special Publication No. 22.

Dendrinos P; Thorpe JP, 1987. Experiments on the artificial regulation of the amino acid and fatty acid contents of food organisms to meet the assessed nutritional requirements of larval, post-larval and juvenile Dover sole (Solea solea (L.)). Aquaculture, 61:121-154.

Dhert P; Sorgeloos P; Devresse B, 1993. Contributions towards a specific DHA enrichment in the live food Brachionus plicatilis and Artemia sp. In: Reinertsen H, Dahle LA, Jørgensen L, Tvinnerheim K, eds. Fish Farming Technology, Balkema, Rotterdam, 109-115.

Dhont J; Lavens P, 1996. Tank production and use of ongrown Artemia. In: Lavens P, Sorgeloos P, eds. Manual on the Production and Use of Live Food for Aquaculture, FAO Fisheries Technical Paper, 361:164-195.

Dhont J; Lavens P; Sorgeloos P, 1991. Development of a lipid enrichment technique for Artemia juveniles produced in an intensive system for use in marine larviculture. In: Lavens P, Sorgeloos P, Jaspers E, Ollevier F, eds. Larvi’91 - Fish & Crustacean Larviculture Symposium, Ostend, Belgium: European Aquaculture Society, Special Publication No. 15, 51-55.

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Estévez A; McEvoy LA; Bell JG; Sargent JR, 1998. Effects of temperature and starvation time on the pattern and rate of loss of essential fatty acids in Artemia nauplii previously enriched using arachidonic acid and eicosapentaenoic acid-rich emulsions. Aquaculture, 165(3/4):295-311; 35 ref.

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Evjemo JO; Coutteau P; Olsen Y; Sorgeloos P, 1997. The stability of docosahexaenoic acid in two Artemia species following enrichment and subsequent starvation. Aquaculture, 155(1/4):135-148; 37 ref.

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Gajardo G; Beardmore JA; Sorgeloos P, 2001. International study on Artemia LXII. Genomic relationships between Artemia franciscana and A. persimilis, inferred from chromocentre numbers. Heredity, 87(2):172-177.

García-Ortega A; Verreth JAJ; Coutteau P; Segner H; Huisman EA; Sorgeloos P, 1998. Biochemical and enzymatic characterization of decapsulated cysts and nauplii of the brine shrimp Artemia at different developmental stages. Aquaculture, 161(1/4):501-514; 32 ref.

Geddes MC; Williams WD, 1987. Comments on Artemia introductions and the need for conservation. In: Sorgeloos P, Bengtson DA, Decleir W, Jaspers E, eds. Artemia Research and its Applications, Vol. 3., Wetteren, Belgium, Universa Press, 19-26.

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Han K; Geurden I; Sorgeloos P, 1999. Enrichment of the nauplii of two Artemia species with docosahexaenoic acid. Crustacean issues, 12:599-604.

Han K; Geurden I; Sorgeloos P, 2000. Comparison of docosahexaenoic acid (22: 6n-3) levels in various Artemia strains during enrichment and subsequent starvation. Journal of the World Aquaculture Society, 31(3):469-475.

Han KyungMin; Geurden I; Sorgeloos P, 2000. Enrichment strategies for Artemia using emulsions providing different levels of n-3 highly unsaturated fatty acids. Aquaculture, 183(3/4):335-347.

Harel M; Ozkizilcik S; Lund E; Behrens P; Place AR, 1999. Enhanced absorption of docosahexaenoic acid (DHA, 22:6n-3) in Artemia nauplii using a dietary combination of DHA-rich phospholipids and DHA-sodium salts. Comparative Biochemistry and Physiology. B, Biochemistry & Molecular Biology, 124(2):169-176; 40 ref.

Hentschel E, 1968. Die postembryonalen Entwicklungsstadien von Artemia salina Leach bei verschiedenen Temperaturen (Anostraca, Crustacea), Zool. Anz, 180:372-384.

Hernandorena A, 1987. An increased dietary tryptophan requirement induced by interference with purine interconversion in Artemia. In: Sorgeloos P, Bengtson DA, Decleir W, Jaspers E, eds. Artemia Research and its Applications, Vol. 2, Universa Press, Wetteren, Belgium, 233-242.

Jones AG; Ewing CM; Melvin MV, 1981. Biotechnology of solar saltfields. Hydrobiologia 82:391-406.

Kellogg VA, 1906. A new Artemia and its life conditions. Science, 24:594-596.

Lavens P, 1989. Intensieve productie en kwaliteitsonderzoek van Artemia adulten en hun nakomelingen (in Dutch). PhD. Ghent, Belgium: Faculty for Agricultural and Applied Biological Sciences, Ghent University.

Lavens P; Baert P; De Meulemeester A; Van Ballaer E; Sorgeloos P, 1986. New developments in the high-density flow-through culturing of brine shrimp Artemia. Journal of the World Mariculture Society, 16:498-508.

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Jean Dhont
Laboratory of Aquaculture & Artemia Reference Center, Faculty Bioscience Engineering, Ghent University, Rozier 44, B - 9000 Ghent, Belgium

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