Artemia (brine shrimp)
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
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 InvasivenessTop of page
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 TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Crustacea
- Class: Branchiopoda
- Order: Anostraca
- Family: Artemiidae
- Genus: Artemia
DescriptionTop 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.
DistributionTop of page
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 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.
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.
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.
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 TableTop of page
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/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|China||Present||Present based on regional distribution.|
|-Fujian||Present||Native||Van Stappen, 2002|
|-Gansu||Present||Native||Van Stappen, 2002|
|-Guangdong||Present||Native||Van Stappen, 2002|
|-Hainan||Present||Native||Van Stappen, 2002|
|-Hubei||Present||Native||Van Stappen, 2002|
|-Jiangsu||Present||Native||Van Stappen, 2002|
|-Liaoning||Present||Native||Van Stappen, 2002|
|-Nei Menggu||Present||Native||Van Stappen, 2002|
|-Ningxia||Present||Native||Van Stappen, 2002|
|-Qinghai||Present||Native||Van Stappen, 2002|
|-Shaanxi||Present||Native||Van Stappen, 2002|
|-Shandong||Present||Native||Van Stappen, 2002|
|-Shanxi||Present||Native||Van Stappen, 2002|
|-Tibet||Present||Native||Van Stappen, 2002|
|-Xinjiang||Present||Native||Van Stappen, 2002|
|-Zhejiang||Present||Native||Van Stappen, 2002|
|India||Present||Native||Invasive||Van Stappen, 2002|
|Indonesia||Present||Introduced||Lavens and Sorgeloos, 2000|
|Iran||Present||Native||Van Stappen, 2002|
|Iraq||Present||Native||Van Stappen, 2002|
|Israel||Present||Native||Van Stappen, 2002|
|Japan||Present||Native||Van Stappen, 2002|
|Kazakhstan||Present||Native||Van Stappen, 2002|
|Korea, Republic of||Present||Native||Van Stappen, 2002|
|Kuwait||Present||Native||Van Stappen, 2002|
|Malaysia||Present||Introduced||Lavens and Sorgeloos, 2000|
|Mongolia||Present||Native||Van Stappen, 2002|
|Myanmar||Present||Introduced||Vanhaecke et al., 1987|
|Pakistan||Present||Native||Van Stappen, 2002|
|Philippines||Present||Introduced||Lavens and Sorgeloos, 2000|
|Saudi Arabia||Present||Native||Van Stappen, 2002|
|Sri Lanka||Present||Native||Van Stappen, 2002|
|Syria||Present||Native||Van Stappen, 2002|
|Taiwan||Present||Native||Van Stappen, 2002|
|Thailand||Present||Introduced||Lavens and Sorgeloos, 2000|
|Turkey||Present||Native||Van Stappen, 2002|
|Turkmenistan||Present||Native||Van Stappen, 2002|
|United Arab Emirates||Present||Native||Van Stappen, 2002|
|Uzbekistan||Present||Native||Van Stappen, 2002|
|Vietnam||Present||Introduced||Lavens and Sorgeloos, 2000|
|Algeria||Present||Native||Van Stappen, 2002|
|Cape Verde||Present||Native||Van Stappen, 2002|
|Egypt||Present||Native||Van Stappen, 2002|
|Eritrea||Present||Native||Lavens and Sorgeloos, 2000|
|Kenya||Present||Native||Van Stappen, 2002|
|Libya||Present||Native||Van Stappen, 2002|
|Madagascar||Present||Native||Van Stappen, 2002|
|Morocco||Present||Native||Amat, 2002; Van Stappen, 2002|
|Mozambique||Present||Native||Van Stappen, 2002|
|Namibia||Present||Native||Van Stappen, 2002|
|Niger||Present||Native||Van Stappen, 2002|
|Senegal||Present||Native||Van Stappen, 2002|
|South Africa||Present||Native||Van Stappen, 2002|
|-Canary Islands||Present||Native||Van Stappen, 2002|
|Tunisia||Present||Native||Van Stappen, 2002|
|Canada||Present||Present based on regional distribution.|
|-Alberta||Present||Native||Van Stappen, 2002|
|-Saskatchewan||Present||Native||Van Stappen, 2002|
|Mexico||Present||Native||Van Stappen, 2002|
|USA||Present||Present based on regional distribution.|
|-Arizona||Present||Native||Van Stappen, 2002|
|-California||Present||Native||Van Stappen, 2002|
|-Hawaii||Present||Native||Van Stappen, 2002|
|-Nebraska||Present||Native||Van Stappen, 2002|
|-Nevada||Present||Native||Van Stappen, 2002|
|-New Mexico||Present||Native||Van Stappen, 2002|
|-North Dakota||Present||Native||Van Stappen, 2002|
|-Oregon||Present||Native||Van Stappen, 2002|
|-Texas||Present||Native||Van Stappen, 2002|
|-Utah||Present||Native||Van Stappen, 2002|
|-Washington||Present||Native||Van Stappen, 2002|
Central America and Caribbean
|Antigua and Barbuda||Present||Native||Van Stappen, 2002|
|Aruba||Present||Native||Van Stappen, 2002|
|Bahamas||Present||Native||Van Stappen, 2002|
|British Virgin Islands||Present||Native||Van Stappen, 2002|
|Costa Rica||Present||Native||Van Stappen, 2002|
|Curaçao||Present||Native||Van Stappen, 2002|
|Dominican Republic||Present||Native||Van Stappen, 2002|
|Haiti||Present||Native||Van Stappen, 2002|
|Jamaica||Present||Native||Van Stappen, 2002|
|Netherlands Antilles||Present||Native||Van Stappen, 2002|
|Nicaragua||Present||Native||Van Stappen, 2002|
|Panama||Present||Introduced||Vanhaecke et al., 1987|
|Puerto Rico||Present||Native||Van Stappen, 2002|
|Saint Kitts and Nevis||Present||Native||Van Stappen, 2002|
|Turks and Caicos Islands||Present||Native||Van Stappen, 2002|
|Argentina||Present||Native||Van Stappen, 2002|
|Bolivia||Present||Native||Van Stappen, 2002|
|Brazil||Present||Present based on regional distribution.|
|-Ceara||Present||Native||Van Stappen, 2002|
|-Rio Grande do Norte||Present||Native||Van Stappen, 2002|
|Chile||Present||Native||Van Stappen, 2002|
|Colombia||Present||Native||Van Stappen, 2002|
|Ecuador||Present||Native||Van Stappen, 2002|
|Peru||Present||Native||Van Stappen, 2002|
|Venezuela||Present||Native||Van Stappen, 2002|
|Bulgaria||Present||Native||Van Stappen, 2002|
|Croatia||Present||Native||Van Stappen, 2002|
|Cyprus||Present||Native||Van Stappen, 2002|
|France||Present||Native||Van Stappen, 2002|
|Greece||Present||Native||Van Stappen, 2002|
|Italy||Present||Native||Van Stappen, 2002|
|Portugal||Present||Native||Van Stappen, 2002|
|Romania||Present||Native||Van Stappen, 2002|
|Russian Federation||Present||Present based on regional distribution.|
|-Eastern Siberia||Present||Native||Van Stappen, 2002|
|-Southern Russia||Present||Native||Van Stappen, 2002|
|-Western Siberia||Present||Native||Van Stappen, 2002|
|Spain||Present||Native||Van Stappen, 2002|
|-Balearic Islands||Present||Native||Van Stappen, 2002|
|Ukraine||Present||Native||Van Stappen, 2002|
|Yugoslavia (Serbia and Montenegro)||Present||Native||Van Stappen, 2002|
|Australia||Present||Present based on regional distribution.|
|-Queensland||Present||Native||Van Stappen, 2002|
|-South Australia||Present||Native||Van Stappen, 2002|
|-Western Australia||Present||Native||Van Stappen, 2002|
|New Zealand||Present||Introduced||Van Stappen, 2002|
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous 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 organisation||Lavens 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)|
ClimateTop of page
|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 TemperatureTop of page
|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 TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|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|
|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 enemiesTop of page
Environmental ImpactTop of page
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 ListTop of page
Animal feed, fodder, forage
- Live feed
ReferencesTop of page
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Laboratory of Aquaculture & Artemia Reference Center, Faculty Bioscience Engineering, Ghent University, Rozier 44, B - 9000 Ghent, Belgium
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