Artemia

Geographical distribution of Artemia View in CoL in Iranian localities

In the present investigation, a diversity of sources including academic research papers, comprehensive checklists, detailed project reports, and conference presentations, alongside firsthand personal observations, were utilized to document the presence and distribution of Artemia within Iran. In table 1 View Table 1 all identified Artemia natural habitats and aquaculture sites are characterized, organizing them by province. This tabulation is enriched with extensive details encompassing geographical coordinates, taxonomic status, and the specific dates of survey or sample collection. Additionally, the geographical distribution of these habitats is illustrated in figure 1 View Fig .

Natural resources

Urmia Lake

Urmia Lake (east of Urmia city, West Azerbaijan Province), was previously ranked as the 20th largest lake on a global scale, with its surface area spanning between 4750 and 6100 km 2 ( Eimanifar & Mohebbi, 2007). The lake's diverse biological and ecological attributes led to its inclusion in the Ramsar Convention on Wetlands in 1975 (https://rsis.ramsar. org/), and it was subsequently designated as one of the 59 world biosphere reserves by UNESCO in 1976 ( Asem et al. 2014, 2016 a). However, over the past twenty years, Urmia Lake has experienced a substantial decrease in its water level, primarily attributed to a series of environmental disturbances and human activities ( Karimi et al., 2016; Golabian, 2011; Asem et al., 2012; Wurtsbaugh et al., 2017; Sima et al., 2021). Post-2007, there has been a marked increase in the lake's salinity, reaching saturation levels exceeding 300 g /l (Asem et al., 2012, 2019). The period between 1995 and 2013 saw the lake losing nearly 60 % of its surface area and over 90 % of its volume ( Schulz et al., 2020). Concurrently, several resource assessments have indicated fluctuations in the population of A. urmiana , with egg densities varying from 339 eggs /liter in 1995, to less than 1 egg /liter between 2008–2012, and then rising to 4.6 eggs/liter during 2018–2020 (see Asem et al., 2012; Ahmadi, 2007; Ahmadi, 2005; Mohebbi et al., 2023 a, b). This environmental crisis has caused a drastic reduction in the annual production of Artemia eggs, decreasing from a minimum of 4243 t dry weight (in the top 0.5 m of water) in 1995 ( Sorgeloos, 1997) to approximately 55 t dry weight (across the entire volume of the lake) in 2018 ( Mohebbi et al., 2023 a). Prior to the drought, the total yield of Artemia eggs, harvested between 2000 and 2002, was estimated to be around 204,000 t (wet weight) ( Mohebbi et al., 2023 a).

Note:

- The type of locality (natural resource/farming site) is not mentioned in this table. For more information, see the text and the original reference(s).

- SUY = survey year; /SAY = sampling year

- non-scientific names were shown in parentheses.

- invalid specific name or invalid description were shown in brackets.

- uncertain geographic coordinates were shown in parentheses.

- The names of localities are in Persian or local languages/accents. Maybe different spellings have been used for a locality in different references

- Phrases of “lake”, “pond”, “catchment”, “Lagoon” and etc. are used with widely explanation in Persian, maybe different phrases have been utilized for one locality in both Persian and English references. Here we followed popular statements in Persian.

- symbol “?” was referred to non-confirmed taxonomic status and unknown locality.

a in some old references “Urmia Lake” also called as “Lake Rezaiyeh”

b non-scientific name

c invalid name

d binominal specific name has mistakenly been written Artemia uromiana .

e binominal specific name has erroneously been written Artemia salinor .

f see Hakupian (1990)

g Artemia parthenogenetica is an invalid binominal specific name, in this table it was replaced to “Parthenogenetic Artemia ” (see Asem et al. 2024b).

h It is also romanized as “Akh Gol” ( Atashbar, 2014). Atashbar et al. (2014) reported Branchinecta orientalis from Agh Gol, but their recorded geographic coordinates (39°33' 18”N 44° 44' 10”E) 1.1 km is out of Agh Gol's shore.

i It is also romanized as “Taschk Lake” ( Löffler, 1959). It is also known as “Nargis Lake” ( Löffler, 1956) and “Nargiz Lake” ( Löffler, 1959). Noori (1995) mistakenly called this lake as “Neyriz Lake”.

j It is also known as “Niris Lake” ( Löffler, 1956) and “Niriz Lake” ( Löffler, 1959).

k It is also romanized as “Maharloo Lake” and “Maharlo Lake”.

l It also romanized as “Schor-gol”, “Schor-Gol” and “Shor Gul”

m This locality might be same as Incheh Lake

n “Anbar Olum Pond” and “Mazraeye Nemuneh” are in Voshmgir District, in Aqqala County.These two sites might be referred to same locality.

o It also known as “ Qom Salt Dome” ( Alvandi & Asil, 2014).

p Aliabad is referred to two localities in Qom Provins   GoogleMaps (34°48'0.79”N 51° 5'11.18”E and 34°17'1.88”N 50°43'20.13”E).

q “Massileh” (mostly romanized as “Masileh”) is the name of plain and its most part is located in Qom Province ( Rahmatizadeh & Jafari, 2014). The exact locality of Artemia site has not been reported.

r “ Markazi Province ” is official term of the province in governmental documents in English ( MOI, 2021). In some references it was erroneously named as “Central Province” (The word Markazi means central in Persian).

s Löffler (1961) called it as “Gavkhaneh”. It is also known as “Gaav Khooni Lake” ( Abatzopoulos et al., 2006a), “Gaav Khooni Wetland” ( Aalamifar et al., 2014) and “Gavkhouni Marsh” ( Goodarzi et al., 2014).

t Abatzopoulos et al. (2006a) and Hafezieh et al. (2015) reported 30°80'E 61°50'N as geographic coordinates, it is referred to Russia (although 80' is also not correct, because maximum amount of each minute is 60 and 30°80'E should be changed to 31°20'E). We changed “N” and “E” following 30°80'N 61°50'E, it is referred to Afghanistan. In   GoogleMaps this table, the geographic coordinates refer to “Varmal Village”.

u Several salty rivers named “Kal-e Shoor” in some provinces ( Abatzopoulos et al., 2006a). Lengths of these rivers are several kilometers and the location of the sampling sites are mostly unclear.

v Banihashemy, per. comm. (2022)

w Abatzopoulos et al. (2006a) and Hafezieh et al. (2015) reported 35°10'E 57°58'N as geographic coordinates, it is referred to Russia. We changed “N” and “E” following 31°20'N 57°58'E, it is referred to Bardaskan County from Shahrabad District in Razavi Khorasan Province   GoogleMaps .

x This site located along “Kal-e Shoor” river.

y It is also known as Nough catchment ( Abatzopoulos et al. 2006a; Hajirostamloo & Pourrabbi 2011; Eimanifar et al., 2014, 2020) and Nough Pool ( Hajirostamloo et al., 2008a). This locality erroneously referred to Yazd Province by Agh et al. (2001).

z Minab has been reported in its original reference (see Agh, 2016). The exact locality should be Tiab (Atashbar per. comm. 2023). Tiab is a village in Tiab Rural District of Minab County.

The initial scientific documentation of Artemia , dating back to the first half of the 10 th century, originates from Urmia Lake and is attributed to the Iranian geographer, Abu-Ishaq Estakhri. In his seminal work, “Al-Masalik wa Al-Mamalik” (commonly known as “Roads and Kingdoms”), Estakhri referred to Artemia as the “aquatic dog”. This codex, penned between 930 and 933 A.D. ( Asem & Eimanifar, 2016; Al-e-Dawoud, 1995). Illustrations in this codex, specifically page 107 depicting a map with Urmia Lake and page 109 detailing the lake and Artemia , are preserved in the Iranian Parliamentary Library.

Subsequent to Estakhri's account, an anonymous Persian geographer made references to Artemia in the manuscript “Hadud Al-alam min Al-Mashriq Ilal-Maghrib” (translated as “The Limits of the World from the East to the West” and known also as “The regions of the world: A Persian geography”) in 982 A.D. (Anonymous, 982). This manuscript describes Artemia as a resilient worm capable of surviving in the saline waters of Urmia Lake (Asem, 2008). Later, in 1275 A.D., another Iranian geographer, Zakariya-Qazwini, reported the existence of an “aquatic dog” as the sole living organism in Urmia Lake in his work, “Asar Al-Beldan wa Akhbar Al-Ebad” (also known as “Monuments of the Land and Histories of the People”) (Zakariya-Qazwini 1275; Asem & Eimanifar, 2016).

Furthermore, in 1877, Muhammad Etemad-Alsaltane, an Iranian politician, documented Artemia in Urmia Lake, describing it as a “herbal-morph animal” in his book “Merat Al-Boldan” (or “Mirror of Cities”) ( Etemad-Alsaltane, 1877). In a later account by George Curzon, a British diplomat, in his 1892 publication “ Persia and the Persian Question”, Artemia from Urmia Lake was identified as a species akin to “Jelly-Fish” ( Curzon, 1892; Sclater, 1893). This series of historical accounts confirms the long-standing recognition and significance of Artemia in the ecological study of Urmia Lake and its surrounding regions.

In the later part of the 19th century, Robert Günther, a British zoologist, received a research grant from the Committee of the Royal Society in the United Kingdom to study the fauna and flora of Urmia Lake ( Günther & Manley, 1899). During the initial stages of his research, in collaboration with his father Albert Günther, who held the position of Keeper of Zoology at the British Museum in London, the Artemia species in Urmia Lake were initially identified as a “Medusa” and a species of “ Branchipus ” ( Günther, 1898 a), and later categorized within “the Artemia group of varieties of the Branchipus type ” ( Günther, 1898 b). However, subsequent investigations led Robert Günther to ascertain that the specimens observed in Urmia Lake were indeed species of the genus Artemia ( Günther, 1899a) . Eventually in 1899, he described Artemia urmiana from Urmia Lake, illustrating a male specimen as well as the brood pouch of a female and the cercopod of both males and females ( Günther, 1899 b).

Notably, there has been some confusion regarding the nomenclature of A. urmiana . In certain references, the binominal specific name A. urmiana has been mistakenly cited to Clark and Bowen (1976), although their work merely referred to the population as “Urmia Lake shrimps” without specifying the name A. urmiana . Additionally, the publication date of A. urmiana has been cited under various years (1890, 1899, and 1900), but it was later clarified that the correct date of description for the species A. urmiana is 1899 ( Asem & Rogers, 2012).

In a significant number of studies spanning from the early 1950s to the late 1980s, Artemia urmiana was mistakenly reported as “ Artemia salina ” (e.g. Plattner, 1955; Löffler, 1956, 1959, 1961; McCarraher, 1972; Daneshgar, 1975; Hoveida, 1976; Saberi, 1978; Kargarnejad, 1986). The name Artemia salina had been generically and inaccurately applied to various species and populations, regardless of their taxonomic status ( Abreu-Grobois & Beardmore, 1982; Gajardo et al., 2002; Asem et al., 2010 a). This misidentification issue has recurred in contemporary studies, particularly those investigating Artemia toxicity.

In 1987, a critical question was raised about the taxonomic status of Urmia Lake Artemia by Barigozzi et al. (1987). Their study focused on the reproductive modes of approximately 500 individuals in a laboratory setting, identifying exclusively parthenogenetic lineages. Through meticulous examination of 24 nauplii to determine chromosome numbers, they recognized three distinct parthenogenetic lineages: diploid (15 specimens), tetraploid (1 specimen), and pentaploid (8 specimens). They proposed that both A. urmiana and parthenogenetic Artemia might exist in Urmia Lake but due to ecological differentiation are located in separate places. Additionally, they concluded that it seems too unlikely that their observation is the result of a replacement of parthenogenetic Artemia with the extinction of A. urmiana ( Barigozzi et al., 1987) . In concurrence with these findings, Azari Takami (1989) reported the observation of both sexual and parthenogenetic Artemia in Urmia Lake, suggesting the coexistence of A. urmiana and parthenogenetic Artemia within the same habitat (see also Barigozzi & Baratelli, 1989; Browne & Bowen, 1991). Further supporting this notion, Ahmadi et al. (1990) documented the coexistence of A. urmiana and parthenogenetic individuals along the western shoreline of the lake, based on samples collected in 1987.

Eskandari (2001) analyzed chromosome numbers and reproductive modes of Artemia from 11 different locations in Urmia Lake. His findings revealed the presence of polyploid parthenogenetic Artemia lineages, although the specific ploidy levels and their distribution, as well as those of A. urmiana , remained unclear. Yar-Mohamadi et al. (2002), in their study conducted between 1998 and 1999, found diploid parthenogenetic Artemia in the Zambil District (west of the lake) and confirmed the presence of both A. urmiana and diploid parthenogenetic Artemia in Urmia Lake.

In a 2005 study, Agh et al. (2007) reported that while only parthenogenetic Artemia were present in three surrounding lagoons, both A. urmiana and parthenogenetic Artemia coexisted within Urmia Lake, which A. urmiana being the dominant population. Further, Asem et al. (2009) observed that parthenogenetic Artemia from Urmia Lake and an adjacent lagoon in Rashakan District were morphologically clustered into two distinct subgroups. Subsequent generations of isolated parthenogenetic Artemia from Urmia Lake exhibited rare males in the laboratory, indicating a diploid lineage ( Asem et al., 2010 b; for more information see Saleem et al., 2019). The results of morphological characteristics of collected Artemia specimens in four stations of Urmia Lake between 2003 and 2004 did not reveal parthenogenetic Artemia ( Asem & Rastegar-Pouyani, 2008 a, b).

Lagoons in the neighborhood of Urmia Lake

In the West Azerbaijan and East Azerbaijan provinces, several lagoons in the neighborhood of Urmia Lake have been identified as natural habitats of Artemia , as described in figure 1 View Fig and table 1 View Table 1 . These lagoons, characterized as diminutive, ephemeral, and shallow bodies of water, occasionally coalesce with the lake during periods of increased precipitation ( Agh et al., 2007; Asem et al., 2009, 2021 b). Comprehensive surveys of these lagoons have exclusively recorded the presence of parthenogenetic Artemia lineages ( Yar-Mohamadi et al., 2002; Hafezieh, 2003 a; Agh et al., 2001; Eimanifar et al., 2007; Asem et al., 2009; Atashbar et al., 2014; Hajirostamloo, 2019; Manaffar et al., 2018). In conclusion, it is inferred that while the lagoons in the vicinity of Urmia Lake harbor parthenogenetic Artemia , the principal body of Urmia Lake   GoogleMaps itself is primarily a habitat for A. urmiana , alongside a limited presence of parthenogenetic Artemia lineage(s). Additionally, unspecified localities within the “Dashte Tabriz” (Tabriz plain) have been recognized as habitats of parthenogenetic Artemia lineages ( Agh, 2016). The geographical expanse of the Tabriz plain, situated between latitudes 37°56' N and 38°17' N, and longitudes 45°30' E and 46°15' E, encompasses approximately 1000 km ², extending from the eastern periphery of Urmia Lake   GoogleMaps to the western boundaries of Tabriz   GoogleMaps city in the East Azerbaijan Province ( Barzegar et al., 2016, 2017; Khatibi & Nadiri, 2021). Therefore   GoogleMaps , the “Lagoons” identified in proximity to Urmia Lake   GoogleMaps in East Azerbaijan, following Agh (2016) checklist, are likely congruent with the Khaslou   GoogleMaps and Sarai   GoogleMaps lagoons enumerated in our catalog ( figure 1 View Fig , table 1 View Table 1 ). Future   GoogleMaps research should prioritize the exploration of Urmia Lake's   GoogleMaps lagoons to augment the existing data on Artemia biodiversity within this region.

Shurabil (Shorabil) Lake

Shurabil Lake (south of Ardabil city, Ardabil Province) is situated at an altitude of approximately 1365 meters above sea level ( Daneshi et al., 2017). Historically, for several decades, this lake was characterized as an alkaline hypersaline water body. In the period of 1974–1975, its salinity was recorded between 107.4 and 121.5 g /l ( Gafurifard et al., 1977). The early documentation of Artemia in Shurabil Lake dates back to 1975, as reported by Pirsamadi (1975). Despite uncertainties in taxonomic classification, this was then referred to as “ A. salina ” ( Pirsamadi, 1975; see also Gafurifard et al., 1977). Since 1995, with the integration of the Balıglı River (Balıglı Chay) and Yamch Dam's spillway in 2003 into the lake's ecosystem, a significant reduction in salinity levels was observed, transitioning the lake from a hypersaline to a freshwater environment ( Sabkara & Makaremi, 2011; Keramati et al., 2016). This ecological shift was accompanied by an expansion of the lake's surface area from 0.74 to 1.7 km 2 (Daneshi, et al., 2017). Consequently, Artemia in Shurabil Lake diminished and eventually became extinct ( Hafezieh, 2003 a; Abatzopoulos et al., 2006 a). In later developments, the narrow-clawed crayfish, Astacus leptodactylus (Crustacea: Decapoda ), was introduced into the lake's ecosystem (see Abd-Almaleki et al., 2009).

Tashk Lake

Tashk Lake (east of Shiraz city, Fars Province) is classified as a saline lake, encompassing an area of roughly 800 km 2 ( Pakzad et al., 2021). The first scientific observation of Artemia species in Tashk Lake was made by Löffler in 1956, who described it as “ A. salina ”, noting that the crustaceans were exclusively female with an average length of 1.13 cm ( Löffler, 1956; see also Löffler, 1959). Further studies have validated that Tashk Lake harbors parthenogenetic Artemia ( Agh et al., 2001; Hafezieh, 2003 a; Abatzopoulos et al., 2006 a). A comprehensive analysis conducted using samples collected in 2011 from Tashk Lake revealed the introduction of the exotic American A. franciscana into this ecosystem ( Shafaie et al., 2012 a, b).

Bakhtegan Lake Bakhtegan Lake (northwest of Neyriz town, Fars Province) alternatively referred to as Neyriz Lake in various scholarly sources ( Löffler, 1956, 1959; Hafezieh, 2003 a), has been identified as a seasonal inland salt lake ( Azareh et al., 2021). In 2003, its surface area was recorded as 1271 km 2 ( Pakzad et al., 2021), with salinity levels ranging between 60 and 250 g /l ( Abatzopoulos et al., 2006 a). During periods of increased rainfall, Bakhtegan Lake is naturally connected to Tashk Lake through a channel ( Hafezieh, 2003 a; Abatzopoulos et al., 2006 a; Safarpour et al., 2018). In some publications, these interconnected lakes are collectively referred to as “Bakhtegan-Tashak Lake” ( Nezami et al., 2013), “Tashak-Bakhtegan Lake” ( Mahabadi et al., 2018), or “Neiriz Lakes”, as listed in the Ramsar Convention on Wetlands (https://rsis.ramsar.org/). The initial reports of Artemia in Bakhtegan Lake date back to Löffler's studies in 1956 and 1959 ( Löffler, 1956, 1959), with subsequent research identifying the crustaceans as parthenogenetic lineage ( Agh et al., 2001; Hafezieh 2003 a; Abatzopoulos et al., 2006 a; Hajirostamloo et al., 2008 a). Additionally, Shafaie et al. (2012 b) documented the presence of the non-native A. franciscana in the lake.

Maharlu Lake

Maharlu Lake (southeast of Shiraz city, Fars Province) is a hypersaline lake with maximum surface area of 258 km 2 ( Samiei et al., 2017) and exhibits a salinity range from 140 to 280 g /l ( Hafezieh, 2003 b). The first occurrence of Artemia in Maharlu Lake was reported by Noori in 1995, although regarding her presented Artemia distribution map, she mistakenly called this lake as “Parishan Lake” (Parishan Lake is a freshwater lake in Jereh and Baladeh District in Fars Province, see Karimian Torghabeh et al., 2021). A field study conducted in 1998 confirmed the presence of parthenogenetic Artemia in Maharlu Lake ( Hafezieh, 2003 b). In subsequent years, numerous studies have corroborated the existence of both parthenogenetic Artemia ( Agh et al., 2001; Abatzopoulos et al., 2006 a; Hajirostamloo et al., 2008 a; Eimanifar et al., 2014; Falahatkar & Rafatnezhad, 2015) and the exotic A. franciscana ( Asem & Aliabadian, 2008; Manaffar et al., 2008; Hoseini & Zare, 2012; Eimanifar et al., 2014, 2020; Manaffar et al., 2018) within the lake. The estimated average annual biomass of Artemia in Maharlu Lake was reported to be 7.54 g /m 3 in June 1998 ( Hafezieh, 2003 b).

Shor Lake

Shor Lake (north of Gorgan city, Golestan Province) encompasses a surface area of 2 km 2 and exhibits a salinity spectrum ranging from 64.1 to 262.4 g /l ( Makhdoumi, 2000). The presence of Artemia in Shor Lake was first documented by McCarraher in 1972, who utilized the conventional nomenclature “ A. salina ” as a species name ( McCarraher, 1972). Subsequent research by Makhdoumi (2000) not only corroborated the existence of Artemia in the lake, it also identified its mode of reproduction as asexual, with the occasional production of male individuals. Makhdoumi (1992) reported that the peak density of Artemia , encompassing various life stages, reached approximately 924.5 individuals/m 3 in the month of April (see also Makhdoumi, 2000). However, due to the influence of the Esteghlal Gorgan Dam's spillway, a notable reduction in the lake's salinity levels was observed, resulting in a transition to freshwater conditions. The latest surveys conducted between 2008 and 2009 indicated that Artemia have subsequently vanished from Shor Lake ( Hami-Tabari, 2011). Agh (2016) reported that the lake has dried up.

Incheh Lake

Incheh Lake (north of Gorgan city, Golestan Province) is situated approximately 20 km from Shor Lake ( Abatzopoulos et al., 2006 a). This lake spans a surface area of 0.6 km 2 and exhibits an annual salinity range between 91.1 and 325.1 g /l ( Makhdoumi, 2000). The first detection of Artemia in Incheh Lake was reported by Makhdoumi in 1992. At its peak, the density of Artemia , including various life stages, was estimated at 2426.7 individuals/m 3 in the month of May ( Makhdoumi, 1992; see also Makhdoumi, 2000). Further investigations by Makhdoumi revealed that Incheh Lake serves as a natural habitat for parthenogenetic Artemia , which also includes a minority group of rare male specimens ( Makhdoumi, 2000).

Mighan Lake

Mighan Lake (northeast of Arak city, Markazi Province) is classified as a shallow, seasonal salt lake with a fluctuating surface area between 81 and 120 km 2 ( Hafezieh et al., 2021). In the year 2019, the lake's average annual salinity was measured at 48.32 g /l ( Hafezieh et al., 2021). Extensive research has consistently demonstrated that Mighan Lake is exclusively inhabited by parthenogenetic Artemia ( Hafezieh, 2003 a; Abatzopoulos et al., 2006 a; Hajirostamloo et al., 2008 a; Manaffar et al., 2015; Eimanifar et al., 2014, 2020; Hesami et al., 2018; Hafezieh et al., 2021). Additionally, in 2019, the total yield of Artemia eggs and biomass in Mighan Lake was quantitatively assessed, revealing figures of 28.875 t and 23.673 t, respectively ( Hafezieh et al., 2021).

Qom Salt Lake

Qom Salt Lake (east of Qom city, Qom Province ) is recognized as the largest seasonal hypersaline lake in Iran ( Safarpour et al., 2018), encompassing an area of 1806 km 2 ( Mirzakani et al., 2017). This lake is also referred to by several names, including “Namak Lake” (with “namak” translating to “salt” in Persian), “Masileh Lake” ( Mostofi, 1971; Mirzakani et al., 2017), and “Aran-Bidgol (Salt) Lake” ( Babavalian et al., 2009; Makhdoumi-Kakhki et al., 2012; Sedghi et al., 2016; Mirzakani et al., 2017; Safarpour et al., 2018; Nasri et al., 2019). Further studies documented the exclusive presence of parthenogenetic Artemia within the lake ( Agh et al., 2001, Hafezieh, 2003 a; Van Stappen, 2002; Mohaghegh & Mohamadpor, 2004; Abatzopoulos et al., 2006 a; Khalili et al., 2007; Hajirostamloo et al., 2008 a; Eimanifar et al., 2014, 2020). Additionally, the area surrounding Qom Salt Lake has been the focus of various temporary pilot studies dedicated to Artemia farming ( Nekoueifard, 2015).

Houze Sultan Lake

Houze Sultan Lake (north of Qom city, Qom Province) is characterized as a seasonal salt lake. Its surface area was recorded at 78.35 km 2 in 2002 and subsequently at 62.65 km 2 in 2013 ( Forootan & Alikhah-Asl, 2019). The lake is alternatively known under several names, “ Qom Lake” (relating to the nearby city south of the lake) “Saveh Lake” (relating to the city to its west) and “Shahi Lake” (with “shahi” translating to “royal” in Persian) ( Shafaroodi, 2009; Forootan & Alikhah-Asl, 2019; Sedghi et al., 2016). The presence of parthenogenetic Artemia in Houze Sultan Lake has been well documented in various scientific reports and studies ( Hafezieh, 2003 a; Mohaghegh & Mohamadpor, 2004; Abatzopoulos et al., 2006 a; Hajirostamloo et al., 2008 a; Soltanifar et al., 2012).

Bazangan Lake

Bazangan Lake (east of Mashhad city, Razavi Khorasan Province) was initially a brackish water body ( Shahnavaz & Ghassemzadeh, 2015). It covered a surface area of 0.69 km 2 and a depth of 13 m ( Adabi & Mohammadzadeh, 1998). The lake is also named as Gol Bibi ( Gholami et al., 2005). Over the past two decades, Batangan Lake has experienced a significant reduction in size, losing about 55 % of its area, attributed primarily to climate change. By 2015, its surface area had diminished to 0.3 km 2, and the salinity levels had increased to 210 g /l ( Mohammadyari et al., 2015). Originally characterized as a hyposaline environment, the lake underwent a transformation into a hypersaline lake. The study conducted by Mohammadyari et al. (2015) reported a parthenogenetic Artemia along with a minor presence of rare male individuals in the lake. The findings led to the conclusion that the colonization of Artemia in Batangan Lake occurred concomitantly with the increase in salinity levels ( Mohammadyari et al., 2015).

Gavkhouni Wetland

Gavkhouni Wetland (southeast of Varzaneh city, Isfahan Province) is also known as Gavkhouni Marsh, with an average surface area of 486 km 2 ( Jaberalansar et al., 2021). This ecological site is enlisted in the Ramsar Convention on Wetlands (https://rsis.ramsar.org/). The occurrence of Artemia in Gavkhouni Wetland was first documented by Löffler in 1961, referred to “ A. saline ” ( Löffler, 1961) . In the summer months, the salt concentration of the wetland's water was recorded at 315 ppt, exhibiting a high magnesium content ( Löffler, 1961). Abatzopoulos et al. (2006 a) reported the presence of only adult female Artemia in the wetland, a finding also supported by Agh et al. (2001). A comprehensive experimental study conducted by Aalamifar et al. (2014) concluded that the samples collected from Gavkhouni Wetland are of a parthenogenetic Artemia .

Hamun-e Jaz Murian Wetland

Hamun-e Jaz Murian Wetland (north of Rameshk city, Kerman Province) is characterized as a seasonal wetland, spanning the geographical region between Kerman and Sistan va Baluchestan. It encompasses an area of approximately 3300 km 2, extending over a length of 100 km and a breadth of 45 km ( Mirzaei & Zibaei, 2019). Observations and studies within this wetland have confirmed the presence of a parthenogenetic Artemia lineage ( Hafezieh, 2003a).

Varmal Lake

Varmal Lake, with a surface area fluctuating between 1.3 and 60 km 2 and a salinity range of 18 to 45 ppt, had been reported from Sistan va Baluchestan Province. The lake's average and maximum depths was between 1.5 and 5 m, respectively ( Mohammad-Piri & Tehrani, 1997). In a study conducted in November 1996, Mohammad-Piri and Tehrani (1997) identified the presence of Artemia in Varmal Lake. Further laboratory investigations confirmed that the Artemia from this lake reproduce asexually ( Mohammad-Piri & Tehrani, 1997).

Surprisingly, the precise geographical location of Varmal Lake   GoogleMaps remains ambiguous. Mohammad-Piri and Tehrani (1997) mentioned that the lake is situated west of Zabol   GoogleMaps city, but did not provide specific coordinates. The village   GoogleMaps of “Varmal” (30°43'32.4" N 61°20'0.8" E), located approximately 35 km southwest of Zabol   GoogleMaps city in the Lutak Rural District of Hamun County, does not have any lake or dried water resource in its vicinity matching the reported surface area. Further   GoogleMaps studies, including those by Abatzopoulos et al. (2006 a) and Hafezieh et al. (2015), have erroneously attributed geographical coordinates to this lake that correspond to a location in Russia. A review of geological and geographical literature yields insufficient information about Varmal Lake. Consequently, the geographic location of “Varmal Lake” is deemed to be an uncertain site in our checklist, warranting further verification and research.

K a l -e S h o o r G o n a b a d

Within the Central Persian Desert basin, a collection of permanent and seasonal saline waterways exists, commonly referred to as “Kal-e Shoor”. In the Razavi Khorasan Province, the Kal-e Shoor Gonabad is a notable example of these saline rivers. After a field investigation focused on Artemia resources in the saline waters of Gonabad during 2001–2002. Ghassemzadeh et al. (2005) reported the presence of parthenogenetic Artemia approximately 35 km north of Gonabad (Banihashemy per. comm. 2022, see also Banihashemy, 2010). The salinity and pH levels in this area ranged from 70.27 to 125.8 g /l and pH 7.5 to 9, respectively, though there are indications that the maximum salinity was alternatively recorded as 155.8 g /l elsewhere in their manuscript. Ghassemzadeh et al. (2005) also identified another habitat of parthenogenetic Artemia in the Yunesi District, situated along the Kal-e Shoor Gonabad River, approximately 15 km northwest of the first site (40 km northwest of Gonabad). This location exhibited salinity levels of 101.8 to 322.9 g /l and a pH range of 7 to 8.6. Artemia densities varied from 101 to 373 individuals/m 3 in Kal-e Shoor and 2 to 190 individuals/m 3 in the Yunesi District ( Ghassemzadeh et al., 2005). Subsequent studies continued to report parthenogenetic Artemia from this river ( Abatzopoulos et al., 2006 a; Banihashemy, 2010), until the discovery of A. franciscana in 2016 ( Agh, 2016).

Additionally, a number of smaller natural habitats, both temporary and seasonal, have been identified in the provinces of Karaj, Lorestan, Kerman, Isfahan, Kohgiluyeh va Boyer Ahmad, and Qom, as summarized in table 1 View Table 1 .

Farming sites

In the context of aquaculture within Iran, Artemia has played a pivotal role as a live feed in larviculture since 1972. This utilization started when Azari Takami pioneered the use of A. urmiana nauplii from Urmia Lake for feeding Persian sturgeon larvae at the Sangar hatchery on the southern coast of the Caspian Sea ( Fayazi, 1994; see also Javanshir, 1986). This period marked a significant increase in the focus on Artemia resources in Iran, emphasizing their application in aquaculture and fisheries.

Kolahi Fishery Farm

At the Kolahi Fishery Farm, an Artemia culture project was initiated in 1993. The farm is located 2 km east of Kolahi village and 30 km southeast of Minab city in the Hormozgan Province. Artemia urmiana was cultivated over a total area of 5100 m 2, within a salinity range of 80 to 110 ppt. Over a seven-month period, Artemia yielded a total biomass of 4,199 kg, translating to an estimated production rate of 1,633 kg per month ( Fayazi, 1994).

Nough Lake

Nough Lake (northwest of Rafsanjan city, Kerman Province) was a natural site that underwent development for fish hatchery establishments in the 1990s. Due to climatic change, notably drought, and a rise in water salinity, promoted Artemia cultivation in the lake by 1999. As of the year 2020, the lake's surface area has expanded from 0.3 km 2 in 1990 to 2 km 2. Annually, Nough Lake yields approximately 20 t of Artemia , both biomass and eggs (www.iranartemia.ir). The project commenced with the introduction of parthenogenetic Artemia ( Abatzopoulos et al., 2006 a; also see Agh et al., 2001), followed by the subsequent colonization of the non-native A. franciscana ( Abatzopoulos et al., 2006 a; Hajirostamloo et al., 2008 b; Hajirostamloo & Pourrabbi, 2011; Eimanifar et al., 2014, 2020). Over the past two decades, reports from the lake have exclusively indicated the presence of A. franciscana ( Hajirostamloo & Pourrabbi, 2011; Eimanifar et al., 2014, 2020).

H e n d i j a n s i t e

In the Hengijan Region of Khuzestan Province, located on the northern coast of the Persian Gulf, a farm was established in the early 2000s with the objective of cultivating Artemia ( Agh et al., 2002) . For this purpose, exotic American A. franciscana , originally sourced from the Mekong Delta in Vietnam, was introduced. Each pond, spanning an area of 112 hectares, yielded a monthly harvest of approximately 15 kg of eggs (wet weight) ( Agh et al., 2002). However, the reported area (each pond with 112 hectares area) does not correspond with the production amounts in this project and pilot studies (see next paragraphs).

Fesendooz site

In 2000, the Iranian Fisheries Science Research Institute established a two-hectare Artemia cultivation site for commercial purposes in the Fesendooz District, located in the southeastern region of Urmia Lake, West Azerbaijan Province. This site was initially populated with A. urmiana sourced from Urmia Lake ( Ahmadi et al., 2012). Subsequent to the filling of certain ponds with saltwater (80 g /l) and fertilization procedures, parthenogenetic Artemia emerged spontaneously, without the introduction of nauplii. Additionally, in other ponds where A. urmiana nauplii were introduced, parthenogenetic Artemia became predominant after a few months (Ahmadi per. comm. 2023). Thus, the Fesendooz District merits recognition as a natural habitat for parthenogenetic Artemia .

Throughout the cultivation period, the site yielded monthly harvests of 32.9 kg dry weight per hectare of parthenogenetic Artemia eggs and 17.3 kg dry weight per hectare of A. urmiana eggs ( Ahmadi et al., 2012). Since the early 2000s, the Fesendooz ponds have been intermittently utilized for Artemia cultivation ( Nekoueifard et al., 2020).

Gobadloo (Ghobadlu) site

Regarding the Gobadloo (Ghobadlu) site, in 2002, an area of 2.3 hectares, divided into eight ponds of 2890 m 2 each, was allocated for the production of A. urmiana and parthenogenetic Artemia in the Gobadloo District, situated in the eastern part of Urmia Lake, East Azerbaijan Province (Atashbar per. comm. 2023; Manaffar et al., 2004; Agh et al., 2007). During the culture period, which lasted 3 to 4 months, the site reported monthly harvests of 36.9 kg dry weight per hectare of parthenogenetic Artemia eggs and 28.8 Kg dry weight per hectare of A. urmiana eggs ( Manaffar et al., 2004). A notable transition was observed in the Gobadloo District pilot project, where the dominance shifted from A. urmiana to parthenogenetic Artemia ( Agh et al., 2007) . Currently, this farming site is not in use for further projects (Atashbar per. comm. 2023).

Shams Abad site

The Shams Abad site, located within the Qanavat Rural District of Qom County in the Qom Province, was developed for the cultivation of both native parthenogenetic Artemia (originating from Qom Salt Lake ) and the non-native A. franciscana . This facility comprised six ponds, collectively covering an area of 600 m 2 (300 m 2 allocated for each treatment), with an average salinity of 50 g /l. A pilot study conducted over 180 days during the summer and autumn of 2013 resulted in the harvesting of 2800 kg of wet biomass of parthenogenetic Artemia and 2200 kg of A. franciscana (wet weight). The study estimated that the average egg production for parthenogenetic Artemia and A. franciscana accounted for approximately 182 and 113 eggs /l, respectively ( Asadpour, 2016 a).

A b S h i r i n s i t e

In the Ab Shirin site, located in the Miyandasht Rural District of Kashan County in the Isfahan Province, a pilot study was conducted to compare the production potential of native parthenogenetic Artemia (sourced from Qom Salt Lake) with that of the exotic American A. franciscana . The site consisted of six ponds, each measuring 50 m 2 and maintaining a salinity of 40 to 50 g /l. The study, which lasted for 105 days during the summer of 2009, revealed that the average number of eggs produced by parthenogenetic Artemia and A. franciscana were 193 and 123.5 egg/l, respectively ( Asadpour, 2016 b).

Additionally, the exotic American A. franciscana has been introduced for economic exploitation in several artificial ponds located along the northern coast of the Persian Gulf and the Sea of Oman, within the provinces of Khouzestan, Bushehr, and Hormozgan ( Jafari & Astani, 2011). Detailed information regarding these introductions is summarized in table 1 View Table 1 , although comprehensive data is not readily available.