Figure 2 - uploaded by Alireza Asem
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Phylogenetic tree of Artemia using COI sequences based on the ML approach. The number behind major nodes denotes bootstrap confidential values. Daphnia tenebrosa (HQ972028) was used as an outgroup. (URM: Artemia urmiana, TIB: Artemia tibetiana, SIN: Artemia sinica, FRA: Artemia franciscana, PER: Artemia prersimilis, SAL: Artemia salina, DI: Diploid parthenogenetic population, TRI: Triploid parthenogenetic population, TETRA: Tetraploid parthenogenetic population, PENTA: Pentaploid parthenogenetic population; abbreviations listed in Table 1).
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Due to the rapid developments in the aquaculture industry, Artemia franciscana, originally an American species, has been introduced to Eurasia, Africa and Australia. In the present study, we used a partial sequence of the mitochondrial DNA Cytochrome Oxidase subunit I (mt-DNA COI) gene and genomic fingerprinting by Inter-Simple Sequence Repeats (IS...
Contexts in source publication
Context 1
... phylogenetic analyses provide evidence that the studied bisexual specimens from three localities of Iran, Nough Catchment (NOG), Mahshar port (MSH) and Maharlu Lake (MAHR), and a locality from Iraq, Garmat Ali (GAA), clustered in the clade of A. franciscana (Figure 2). In addition, all parthenogenetic populations clustered in two separated clades that shared a common ancestor with urmiana. ...
Context 2
... et al. [2] have documented that di-and triploid parthenogenetic brine shrimps are maternally related to A. urmiana, while tetra-and pentaploid lineages shared a common maternal ancestor with A. sinica. Based on the mitochondrial COI dataset, all examined parthenogenetic individuals have grouped in a close evolutionary relationship with A. urmiana (Figure 2). Our results have demonstrated that they should include di-and/or triploids. ...
Context 3
... observation has also been confirmed by the phylogenetic tree. Although individuals of eight Artemia populations have been exactly located in sub-clades of diand/or triploid, eight out of nine specimens of CAM from Turkey have been placed in a particular clade (P2), in a close connection with A. urmiana (Figure 2). This finding has also been confirmed by haplotype distribution (Figure 3). ...
Citations
... Asem et al. (2021) determined the evolutionary relationship and the genetic variation of bisexual and parthenogenetic Artemia using three mitochondrial and two nuclear markers, they find that the diploid parthenogenetic Artemia are closely related to Artemia urmiana and tetraploids share a common ancestor with Artemia sinica. With time, an unusually high degree of divergence has produced among populations because of considerable changes in environmental conditions, such as salinity and temperature which consequently generated variability in reproductive, lifespan and physiological traits, population size (Browne 1992;Eimanifar et al. 2020). Parthenogenetic populations have been predicted to respond less to environmental change which was observed in our study for reproduction and survival of the Bethioua population. ...
... The parthenogenetic Artemia populations from Portugal showed vulnerability and great variability in the physiological response to different abiotic conditions, suggesting possible local adaptations in response to different selective pressures experienced like other Artemia parthenogenetica populations (Pinto et al. 2013). The environmental parameters, such as salinity and temperature, play an important Artemia salina (S), + + : present study, + : population studied by Ghomari (2013) role in genetic structure and population size during evolution of local Artemia populations (Eimanifar et al. 2020). ...
Morphological and reproductive characteristics of brine shrimp Artemia from two western Algerian populations (El Melah and Timimoun) were studied for the first time alongside the northwestern population (Bethioua). Sexual, survival, growth, and reproduction were recorded for each population raised under standardized culture conditions. The morphological results showed that Artemia from the Timimoun and El Melah populations are bisexual, while the Bethioua population is parthenogenetic. After 20 days at salinity of 80 PSU, higher survival rates were found for the El Melah (85.5%) and Timimoun (72.0%) populations, while the Bethouia population showed the lowest survival rate (30.2%). Total body lengths of naupliar stages of the parthenogenetic populations were significantly longer than those of the bisexual populations. The Bethouia population displayed the highest absolute growth rate resulting in the largest adults compared to the bisexual populations. El Melah and Timimoun females were significantly larger morphologically in seven of the nine morphological characteristics compared to their respective males. The Bethouia females had smaller total and abdominal lengths compared to the females from the bisexual populations. The bisexual strains were significantly different from the parthenogenetic population in 9 of the 11 reproductive characteristics. Bethouia females, however, had the highest offspring/day/female and longest post-reproductive period compared to bisexual females. The Timimoun population adapted reproductive strategies differently compared to the other two populations: a late maturity (21.6 days), and more offspring/females (65.1). The data from the study will help future management and potential development of the Algerian brine shrimp populations.
... The other four species are native to the Old World: A. salina (Linnaeus, 1758) in the Mediterranean region, A. sinica Cai, 1989in China, A. urmiana Günther, 1899 in Lake Urmia (Iran) and the Crimean Peninsula, and A. tibetiana Abatzopoulos, Zhang & Sorgeloos, 1998, in the Tibetan or Qinghai-Tibet Plateau. Artemia franciscana has been anthropogenically introduced in Eurasia and Australia (Zheng et al., 2004;Amat et al., 2005;Abatzopoulos et al., 2009;Scalone & Rabet, 2013;Eimanifar et al., 2014Eimanifar et al., , 2020Asem et al., 2018Asem et al., , 2021aSaji et al., 2019;Shen et al., 2021). Obligate parthenogenetic linegaes widely distributed in Eurasia, Africa, and Australia consist of different ploidy (di-, tri-, tetra-, penta-and heteroploids) (Sun et al., 1999;Asem & Sun 2014a, b;. ...
... The primary complication of the submitted COI sequence of A. frameshifta (LC195588) is the presence of several stop codons (indicating that the sequence is a NUMT), which in this case, renders the sequence used unreliable. Artemia frameshifta is therefore a nomen dubium (Asem et al., 2018(Asem et al., , 2022Eimanifar et al., 2020). Further studies show that the Mongolian populations resemble A. sinica (unpublished data;S. ...
... Previous phylogenetic analyses based on the mitochondrial COI and the nuclear ITS1 marker considered the American A. franciscana and Asian Artemia as sister clades, with A. franciscana as a basal for this group. Additionally, A. sinica was reported as the basal clade for Asian Artemia (Baxevanis et al., 2006;Kappas et al., 2009;Maniatsi et al., 2011;Eimanifar et al., 2014Eimanifar et al., , 2020Saji et al., 2019;Asem et al., 2018Asem et al., , 2021a. Our findings, using the complete mitochondrial sequences, also suggest that A. sinica and the American A. franciscana would be divided with a basal node ("ancestral node," according to Omland et al., 2008), in which A. sinica is basal to other Asian species of Artemia. ...
Species of Artemia are regionally endemic branchiopod crustaceans composed of sexual species and parthenogenetic lineages, and represent an excellent model for studying adaptation and speciation to extreme and heterogeneous hypersaline environments. We tested hypotheses of whether populations from the Tibetan Plateau belong to A. tibetiana Abatzopoulos, Zhang & Sorgeloos,1998 and whether a population from Kazakhstan is a new species, using other Asian species of Artemia as outgroups. We conducted a multitrait phylogenetic study based on the complete mitogenome, mitochondrial (COI, 12S, 16S) and nuclear (microsatellites, ITS1) markers, and a suit of uni-and multivariate morphological traits. Our results led to the discovery of two new species, one from the Tibetan Plateau (Haiyan Lake) in China (Artemia sorgeloosi n. sp.) and a second from Kazakhstan (Artemia amati n. sp.). Our analysis demonstrate that A. tibetiana and A. amati n. sp. are monophyletic, whereas A. sorgeloosi n. sp., and A. tibetiana are polyphyletic. Evolutionary relationships based on mitochondrial and nSSR markers suggest that A. tibetiana may have arisen from a past hybridization event of a maternal ancestor of A. tibetiana with A. sorgeloosi n. sp. or its ancestor. We present the complete mitogenome of A. tibetiana, A. amati n. sp., and A. sorgeloosi n. sp. We also provide a novel taxonomic identification key based on morphology, emphasizing the phenotype as a necessary component of the species concept.
... Four bisexual species are native to the Old World namely Artemia salina (Linnaeus, 1758), Artemia urmiana Günther, 1899, Artemia sinica Cai, 1989, and Artemia tibetiana Abatzopoulos et al. (1998). The other three bisexual species are located in the New World consisting of Artemia monica Verrill, 1869, Artemia franciscana Kellogg, 1906, and Artemia persimilis Piccinelli and Prosdocimi, 1968 [18,20,21]. Obligate parthenogenetic Artemia taxa have di-, tri-, tetra-and pentaploid populations [19]. ...
In the previously published mitochondrial genome sequence of Artemia urmiana (NC_021382
[JQ975176]), the taxonomic status of the examined Artemia had not been determined, due to partheno�genetic populations coexisting with A. urmiana in Urmia Lake. Additionally, NC_021382 [JQ975176]
has been obtained with pooled cysts of Artemia (0.25 g cysts consists of 20,000–25,000 cysts), not a
single specimen. With regard to coexisting populations in Urmia Lake, and intra- and inter-specific
variations in the pooled samples, NC_021382 [JQ975176] cannot be recommended as a valid se�quence and any attempt to attribute it to A. urmiana or a parthenogenetic population is unreasonable.
With the aid of next-generation sequencing methods, we characterized and assembled a complete
mitochondrial genome of A. urmiana with defined taxonomic status. Our results reveal that in
the previously published mitogenome (NC_021382 [JQ975176]), tRNA-Phe has been erroneously
attributed to the heavy strand but it is encoded in the light strand. There was a major problem in the
position of the ND5. It was extended over the tRNA-Phe, which is biologically incorrect. We have
also identified a partial nucleotide sequence of 311 bp that was probably erroneously duplicated
in the assembly of the control region of NC_021382 [JQ975176], which enlarges the control region
length by 16%. This partial sequence could not be recognized in our assembled mitogenome as well
as in 48 further examined specimens of A. urmiana. Although, only COX1 and 16S genes have been
widely used for phylogenetic studies in Artemia, our findings reveal substantial differences in the
nucleotide composition of some other genes (including ATP8, ATP6, ND3, ND6, ND1 and COX3)
among Artemia species. It is suggested that these markers should be included in future phylogenetic
studies.
... Unintentional escapes caused by normal use in hatcheries and/or transmission by migratory waterfowl should be considered as a secondary factor in the distribution of A. franciscana in new habitats. At present, A. franciscana has been colonized in numerous regions across Eurasia, especially in the Mediterranean (Amat et al., 2005;Mura et al., 2006;Van Stappen, 2008;Muñoz, 2009;Ben Naceur et al., 2010, Eimanifar et al., 2014Scalone and Rabet, 2013;Horvath et al., 2018;Saji et al., 2019;Eimanifar et al., 2020) and Australia (Asem et al., 2018). ...
Artemia franciscana, native to America, has recently colonized as non-indigenous population in Asia, Europe, North Africa, and Australia. We evaluated the effects of the colonization of A. franciscana on genetic differentiation in new environments in the United Arab Emirates (UAE). We used the COI marker to determine the genetic structure and origins of exotic populations in the UAE. Results confirmed the colonization of A. franciscana in two localities. Invasive populations of A. franciscana had significantly lower genetic variation than native populations in the Great Salt Lake and San Francisco Bay. Results showed that the studied populations could not have colonized directly from natural American habitats, and they possibly were from secondary introduction events of other non-indigenous populations. Genetic analysis yielded different demographic patterns for the studied invasive populations. The population in Al Wathba Wetland Reserve (AWWR) demonstrated demographic expansion, whereas in Godolphin Lakes (GL), it reached a demographic equilibrium. Neutrality tests showed an excess of recent and historical mutations in the COI gene pool of invasive AWWR Artemia in the new environment. The results suggest that different ecological conditions in new environments can exert selective pressures during the introduction of an exotic population, which can affect genetic variation.
... Unintentional escapes caused by normal use in hatcheries and/or transmission by migratory waterfowl should be considered as a secondary factor in the distribution of A. franciscana in new habitats. At present, A. franciscana has been colonized in numerous regions across Eurasia, especially in the Mediterranean (Amat et al., 2005;Mura et al., 2006;Van Stappen, 2008;Muñoz, 2009;Ben Naceur et al., 2010, Eimanifar et al., 2014Scalone and Rabet, 2013;Horvath et al., 2018;Saji et al., 2019;Eimanifar et al., 2020) and Australia (Asem et al., 2018). ...
Artemia franciscana, native to America, has recently colonized as non-indigenous population in Asia, Europe, North Africa, and Australia. We evaluated the effects of the colonization of A. franciscana on genetic differentiation in new environments in the United Arab Emirates (UAE). We used the COI marker to determine the genetic structure and origins of exotic populations in the UAE. Results confirmed the colonization of A. franciscana in two localities. Invasive populations of A. franciscana had significantly lower genetic variation than native populations in the Great Salt Lake and San Francisco Bay. Results showed that the studied populations could not have colonized directly from natural American habitats, and they possibly were from secondary introduction events of other non-indigenous populations. Genetic analysis yielded different demographic patterns for the studied invasive populations. The population in Al Wathba Wetland Reserve (AWWR) demonstrated demographic expansion, whereas in Godolphin Lakes (GL), it reached a demographic equilibrium. Neutrality tests showed an excess of recent and historical mutations in the COI gene pool of invasive AWWR Artemia in the new environment. The results suggest that different ecological conditions in new environments can exert selective pressures during the introduction of an exotic population, which can affect genetic variation.
... Four bisexual species are native to the Old World namely Artemia salina (Linnaeus, 1758), Artemia urmiana Günther, 1899, Artemia sinica Cai, 1989, and Artemia tibetiana Abatzopoulos et al. (1998). The other three bisexual species are located in the New World consisting of Artemia monica Verrill, 1869, Artemia franciscana Kellogg, 1906, and Artemia persimilis Piccinelli and Prosdocimi, 1968 [18,20,21]. Obligate parthenogenetic Artemia taxa have di-, tri-, tetra-and pentaploid populations [19]. ...
In the previously published mitochondrial genome sequence of Artemia urmiana (NC_021382 [JQ975176]), the taxonomic status of the examined Artemia had not been determined, due to parthenogenetic populations coexisting with A. urmiana in Urmia Lake. Additionally, NC_021382 [JQ975176] has been obtained with pooled cysts of Artemia (0.25 g cysts consists of 20,000–25,000 cysts), not a single specimen. With regard to coexisting populations in Urmia Lake, and intra- and inter-specific variations in the pooled samples, NC_021382 [JQ975176] cannot be recommended as a valid sequence and any attempt to attribute it to A. urmiana or a parthenogenetic population is unreasonable. With the aid of next-generation sequencing methods, we characterized and assembled a complete mitochondrial genome of A. urmiana with defined taxonomic status. Our results reveal that in the previously published mitogenome (NC_021382 [JQ975176]), tRNA-Phe has been erroneously attributed to the heavy strand but it is encoded in the light strand. There was a major problem in the position of the ND5. It was extended over the tRNA-Phe, which is biologically incorrect. We have also identified a partial nucleotide sequence of 311 bp that was probably erroneously duplicated in the assembly of the control region of NC_021382 [JQ975176], which enlarges the control region length by 16%. This partial sequence could not be recognized in our assembled mitogenome as well as in 48 further examined specimens of A. urmiana. Although, only COX1 and 16S genes have been widely used for phylogenetic studies in Artemia, our findings reveal substantial differences in the nucleotide composition of some other genes (including ATP8, ATP6, ND3, ND6, ND1 and COX3) among Artemia species. It is suggested that these markers should be included in future phylogenetic studies.
Artemia franciscana is a universal live feed in aquaculture, and it has been reported as an invasive species in many Asian hypersaline ecosystems. The present observations illustrated the pre-and postembryonic development stages of the A. franciscana population confined to the Indian saltern of Kelambakkam. We observed their growth patterns during various hydration periods with specific time intervals. Results showed differences in the development stages with respect to unique identity. Interestingly, a period of hydration showed notable cellular movement toward clockwise positions in the hydrating cysts. After 10 h of hydration, blastocoel appeared, accelerating the dynamic route of nuclei movement. At the end of the invagination, the embryo burst out of the cyst, and a sequence of emerging stages was noted. With reference to light microscopic observations, a series of developmental stages were observed, and each instar was documented by developing limb buds of nauplii. Excitingly, the 10th and 11th instar stages reveal sexual differentiation between male and female individuals. Thus, the laboratory culture study clearly documented the different developmental stages with their specific characteristic features. However, further molecular study would provide a cellular basis for understanding the early development of A. franciscana. K E Y W O R D S aquaculture, artemia, cyst, embryonic development, histology
Hypersaline lakes in arid and semi-arid areas are unique ecosystems that harbor unique extremophile organisms, such as Artemia, the paradigmatic example of adaptation to harsh living conditions. We assessed the mitogenomic biodiversity of Artemia species from the Tibetan Plateau, China, a remote and yet minimally disturbed ecosystem with a variety of hypersaline lakes. Analysis of ten Tibetan salt lakes demonstrated the occurrence of two regionally endemic species, A. tibetiana and A. sorgeloosi, with the latter being the dominant species with eight localities. Both species coexist in Jingyu and Jibu lakes, representing the first case of natural distribution overlap between sexual Artemia species. Artemia sorgeloosi exhibits higher genetic diversity and interpopulation differences, a result consistent with the heterogeneity of local salt lakes, local Artemia population demographics, and their adaptive potentials. Significant FST values demonstrate a gene flow barrier between A. sorgeloosi populations that is compatible with an “island biogeography” distribution pattern, making the Tibetan Plateau a sort of natural laboratory to study intraspecific population differences. Artemia sorgeloosi and the exotic A. franciscana were found coexisting in Yangnapen Lake, demonstrating the ability of this invasive species to colonize high-altitude inland habitats, and the need to monitor its presence and eventual expansion.
Artemia is the most common live food that is used in aquaculture worldwide. This study reportson biometrical variation in the cysts of the introduced, originally American Artemia franciscanafrom 24 non-native localities and two native habitats, in Asia and the U.S.A., respectively. Theresults show, that the largest diameter of untreated cysts, the largest diameter of decapsulatedcysts, and a thicker chorion usually are found in invasive populations. Because of the small cysts,which have an effect on an increasing quantity per unit weight and thus could be the cause ofincreased hatching efficiency, commercial productions of A. franciscana cysts from native sourcesshould potentially be considered higher quality than productions from non-indigenous environments.Principal Component Analysis revealed that all cyst batches from San Francisco Bay were classifiedin one group and most of the invasive populations could be arranged in another, separate group.Although the diameter of the decapsulated cyst and chorion thickness showed a negative andsignificant correlation among invasive populations, there was no significant relationship within nativepopulations. These observations contrast with biometrical patterns of parthenogenetic populations.
