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Observed mismatch distributions and their curve fit to simulated models of demographic expansion. MUL, Mulgundawa; SKI, St Kilda; HED, Port Hedland; DAM, Dampier; GSL, Great Salt Lake; SFB, San Francisco Bay.

Observed mismatch distributions and their curve fit to simulated models of demographic expansion. MUL, Mulgundawa; SKI, St Kilda; HED, Port Hedland; DAM, Dampier; GSL, Great Salt Lake; SFB, San Francisco Bay.

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Article
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Native American Artemia franciscana has become an introduced species in the Old World due to the rapid development of the aquaculture industry in Eurasia. The recent colonisation of A. franciscana in Mediterranean regions and Asia has been well documented, but Australia is a continent where the dispersal of this species is not well understood. In t...

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... native populations of A. franciscana. These revealed that the SKI, HED and GSL localities had a unimodal pattern, whereas MUL and SFB localities showed a pattern likely to be multimodal. Additionally, the indices of SSD and Hri for all examined localities were non-significant, except for MUL, where we observed a significant SSD value (P < 0.001) (Fig. ...
Context 2
... native populations of A. franciscana. These revealed that the SKI, HED and GSL localities had a unimodal pattern, whereas MUL and SFB localities showed a pattern likely to be multimodal. Additionally, the indices of SSD and Hri for all examined localities were non-significant, except for MUL, where we observed a significant SSD value (P < 0.001) (Fig. ...

Citations

... The introduction of A. franciscana around the world represents a risk for local endemic Artemia populations and species; thus, there is a need to assess and protect their biodiversity at the gene, population, and species levels , 2021, Laikre et al. 2020; Thomson et al. 2021, Hvilsom et al. 2022. Artemia franciscana has tremendous intraspecific ecological plasticity (Gajardo et al. 2002, Gajardo and Beardmore 2012, and has invaded Old World and Australian habitats, sometimes displacing local species (Zheng et al. 2004, Scalone and Rabet 2013, 2020, Asem et al. 2018, 2021, Horváth et al. 2018, Saji et al. 2019, Shen et al. 2021, Wu et al. 2022. Its ecological plasticity allows it to tolerate a variety of salinity concentrations (7 to 340 g/L), ionic composition (chloride and sulphate/carbonate), altitude (sea level to 4500 m a.s.l.), and climate (tropical to subarctic) (Post and Youssef 1977, Zheng 1997, Van Stappen et al. 2003, Asem et al. 2012, 2019, Gajardo and Beardmore 2012, Zheng and Sun 2013. ...
... Due to its adaptive capabilities and high reproductive potential, it was introduced in new regions around the world for economic exploitation; but, it outcompetes most native Old World populations (Amat et al. 2005, Mura et Shen et al. 2021, Wu et al. 2022. It has also been introduced to Australia ( Asem et al. 2018). However, the limited amount of A. franciscana in Yangnapen Lake (< 8% of Artemia community composition) likely reveals an initial dispersal event but has not yet become established at this site. ...
... Regarding mismatch distribution, few populations present a multimodal pattern. This finding suggests demographic equilibrium (see Ferreira et al. 2017, Asem et al. 2018, 2021, Santos et al. 2018, Srinivas et al. 2020, Ualiyeva et al. 2022) which can be supported by nonsignificant Tajima's D test values. The present study demonstrates a negative and significant Tajima's D value for most populations/markers with unimodal patterns of mismatch distribution including both species of A. tibetiana and A. sorgeloosi. ...
Article
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.
... 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. ...
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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.
... It would be more appropriate to consider the risks of introduction to the Antarctic of a competing alien species for B. gaini. An example from another part of the world is an invasive anostracan species of the genus Artemia, native to North America that currently threatens the Mediterranean populations of this genus (Asem et al., 2018). Many invertebrates, especially those associated with astatic water bodies, such as B. gaini, have dormant forms or cysts, which increases the risk of their transmission on footwear over long distances. ...
... Artemia is produced in every continent except for Antarctica [42][43][44][45][46][47]. Moreover, previous studies have also shown some information about the bibliometric knowledge on live feed such as Artemia [47], but did not focus on a detailed explanation of the studies, in contrast to the present study. ...
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An increasing number of scientists since 1970 has examined Artemia as an important species in aquaculture-related fields. However, a global scientometric review of Artemia literature is still lacking, which is the objective of this research. Using a CiteSpace analysis, the distribution of core authors and institutions, highly cited keywords and papers, author and journal contributions, and hot topics in the literature, as well as a co-citation analysis, particularly regarding authors, journals, documents, and clusters, were determined. Hence, 8741 relevant publications were generated from the Web of Science Core Collection database. The results revealed that the most significant contributions in Artemia research primarily originated from the USA, Brazil, Spain, India, China, and Belgium. Moreover, Artemia research focused mainly on top keywords such as brine shrimp and antimicrobial activity. Emerging trends related to Artemia research were Atlantic halibut, elongation factor, Artemia salina, lean protein, inert diet, alpha-crystallin protein, and Artemia embryo. At the same time, the study generated a vast total of 45 co-citation clusters. The present study provides the existing body of knowledge on Artemia research by sharing a visual knowledge map. This study offers a valuable perspective and profound understanding for researchers, farmers, and consortia interested in promoting Artemia as a sustainable live food in the global aquaculture industry.
... Many works have been devoted to the study of the molecular genetic diversity of Artemia in different regions [5,18,19]; however, there are still some practically unexplored regions. One of these is Crimea, the largest peninsula in the Black Sea (27,000 km 2 ). ...
... Once in a new region, A. monica begins to change rapidly, adapting to the conditions of the new region [18,19,51,52]. The rapid variability and adaptability of A. monica under new conditions are facilitated by the fact that the species has different alternative gene expression patterns [16,53]. ...
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Many works have been devoted to the study of the molecular genetic diversity of Artemia in different regions; however, there are regions such as Crimea, the largest peninsula in the Black Sea, which has seen few studies. Artemia specimens from several Crimean hypersaline lakes were analyzed using the mitochondrial marker cytochrome oxidase C (COI). The analyzed individuals from bisexual populations formed clades with the species A. salina, A. urmiana, A. sinica, and A. monica (=A. franciscana). A. sinica and A. monica had not been recorded in Crimea previously. In Lake Adzhigol, the three species A. urmiana, A. sinica, and A. monica were found at the same time, which has not been noted anywhere before. In the Crimean lakes, a total of 10 haplotypes were found, six of them for the first time: Once for A. monica, once for A. sinica, and four for A. salina. Those haplotypes may be regarded as endemic to Crimea. In the 1990s, experiments were carried out in Lake Yanyshskoe using mainly purchased cysts of Artemia, so A. monica and A. sinica were introduced into Crimea and could then have easily been spread by birds to other Crimean lakes.
... Samples L. Sainz-Escudero et al. Additionally, to our data, all the available Artemia cox1 sequences available in GenBank (Valsala et al. 2005;Hou et al. 2006;Tizol-Correa et al. 2009;Muñoz et al. 2008Muñoz et al. , 2010Muñoz et al. , 2013Maniatsi et al. 2009Maniatsi et al. , 2011Maccari et al. 2013b;Eimanifar and Wink 2013;Eimanifar et al. 2014Eimanifar et al. , 2015Eimanifar et al. , 2016Asem et al. 2016Asem et al. , 2019Asem et al. , 2020Naganawa and Mura 2017;Horváth et al. 2018) and one of Branchinecta ferox used as outgroup (LT821334 [Rodríguez-Flores et al. 2017]) were retrieved in order to build a dataset represented by 1505 sequences, that allowed us to depict the structuring of the genus through the Neighbour Joining analysis and to perform phylogeographic analyses. Some dissimilar sequences that featured stop codons when traduced to amynoacids were removed from the analyses due to the existence of pseudogenes according to Rode et al. (2021). ...
Article
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Brine shrimps ( Artemia ) have undergone geographic range and demographic expansions as a result of their interaction with humans since the beginning of salt harvesting. This interaction has favoured the expansion of some species but compromising the survival of others. Mediterranean native populations of Artemia salina from coastal salterns and lagoons are facing the presence and expansion of the introduced and invasive American species Artemia monica (= A . franciscana ). However, this species could not be the only threat. Parthenogenetic populations of the Asian species A . urmiana and A . sinica are widespread along the Mediterranean and other areas of the world. In this work, with the use of large cox1 and mitogenomic datasets, phylogenetic and phylogeographic inferences, and a time calibrated tree, we confirmed the Asian origin and recent arrival of the current Western Mediterranean parthenogenetic populations of Artemia . In addition, the replacement of Iberian populations of A . salina by Asiatic parthenogenetic populations lead us to recognize parthenogens as invasive. Current salterns development and commercial importance of Artemia make human-mediated introduction probable. These results demonstrate again the impact that changing human interests have on population expansion or decline of species adapted to anthropogenic habitats. Artemia salina decline makes urgent the implementation of conservation measures such as its use in fish farming and salt production or its inoculation in inland salterns.
... The genus Artemia comprises seven (Rogers 2013) or possibly nine (Naganawa and Mura 2017) species. Although Artemia is widespread and common in hypersaline lakes in most continents, only Artemia franciscana Kellogg and Artemia parthenogenetica Bowen & Sterling are present in Australia (McMaster et al. 2007;Asem et al. 2018). A. franciscana was introduced to Australia by humans to aid in salt production and is overwhelmingly restricted to constructed evaporative ponds (salt works; Timms and Hudson 2009;Asem et al. 2018). ...
... Although Artemia is widespread and common in hypersaline lakes in most continents, only Artemia franciscana Kellogg and Artemia parthenogenetica Bowen & Sterling are present in Australia (McMaster et al. 2007;Asem et al. 2018). A. franciscana was introduced to Australia by humans to aid in salt production and is overwhelmingly restricted to constructed evaporative ponds (salt works; Timms and Hudson 2009;Asem et al. 2018). A. parthenogenetica may have also been introduced into salt works by humans (McMaster et al. 2007), but its presence in a range of lakes in south-western Australia could be the result of intercontinental bird-mediated dispersal followed by local dispersal (McMaster et al. 2007). ...
... All Artemia species, including the two species found in Australia, have high dispersal capacity because their cysts float and are effectively transported by animals and wind (Timms and Hudson 2009). Despite this, A. franciscana does not seem to be spreading (Timms and Hudson 2009;Asem et al. 2018), but the distribution of A. parthenogenetica is increasing in southwestern Australia, where it is mostly colonising degraded salt lakes (McMaster et al. 2007). It is unclear how the further spread of A. parthenogenetica may affect Parartemia species, but currently A. parthenogenetica appears limited to lakes not already occupied by Parartemia (McMaster et al. 2007). ...
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This study synthesises information on the biology of the unique and diverse halophilic macroinvertebrates of Australian salt lakes, focusing on gastropods and crustaceans. This information is needed to evaluate and manage the threats posed to these invertebrates by increased periods of drought and secondary salinisation. Most of these species are endemic to Australian salt lakes, and some have adapted to extreme conditions (e.g. salinities >100gL<sup>–1</sup> and pH <5). This study identifies key general findings regarding the taxonomy, ecology and life histories of these invertebrates, such as that many ‘new’ species have been uncovered in the past 20 years, with more likely to come. The study also identifies critical knowledge gaps, such as the need to elucidate the abiotic and biological drivers of the field distributions of species, including why some species are widespread and common whereas other congeneric species are rare or have narrow distributions. Those species that are either restricted to low salinity environments or survive dry periods as aestivating adults (as opposed to desiccation-resistant eggs) are probably the most vulnerable to increasing salinisation and drought. Future work should prioritise the development of a sound taxonomy for all groups, because this is needed to underpin all other biological research.
... Eventually, this had led to the invasion of A. franciscana into various countries (Muñoz et al., 2009;Ruebhart et al., 2008) including in Asia, Africa, and Europe, resulting in the elimination of native Artemia species (Ben Naceur et al., 2010;Scalone and Rabet, 2013;Muñoz et al., 2014;Amat et al., 2007). With the evidence of morphology, morphometric and a mitochondrial DNA (COXI) marker, recent studies confirm the invasion of A. franciscana in Egyptian and Australian hypersaline habitats (Sheir et al., 2018;Asem et al., 2020). Similarly, invasion of A. franciscana and replacement of the native population of Artemia parthenogenetica was confirmed in Indian salterns by using ITS-I gene sequences (Vikas et al., 2012). ...
... KBM haplotypes never shared their genetic character with other sampling sites while VDM, TUT, and NGC showed strong genetic dispersal between population. This observation can be supported by an earlier report on invasion and occurrence of A. franciscana from Australian salterns demonstrated by the COX1 gene (Hd = 0.533) (Asem et al., 2020). The deep-sea decapod crustacean species Aristeus antennatus also showed a high level of haplotype diversity based on the mtDNA control region analysis (H = 0.884 to 0.989) (Maggio et al., 2009;Eimanifar et al., 2014). ...
Article
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The invader brine shrimp Artemia franciscana is a micro-crustacean with diverse morphology and morphometric traits that are found in hypersaline habitats of Asian countries. In this study, we used the mtDNA control region to assess the impact of colonization of A. franciscana on genetic diversity and population structure encountered in different sampling sites of the southeast coast of India. Samples were collected from four hypersaline habitats includes Kelambakkam (KBM); Vedaranyam (VDM); Tuticorin (TUT) and Nagarcoil (NGC). Genetic diversity of the populations were assessed based on the sequence of a conserved region of the mtDNA control region (CR). For this study, a primer based on the conserved region of a closely related crustacean group was used to sequence the 493 bp of the mtDNA CR. Results indicated a clear demarcation with the existence of sequence divergence between the populations studied. Maximum likelihood and Bayesian phylogenetic studies revealed that the population of TUT, VDM and NGC formed a single cluster whereas the KBM formed a separate cluster with distinct genetic characteristics. The mean haplotype diversity and nucleotide diversity of the four populations were found to be 0.742 ± 0.060 and 0.012 ± 0.016, respectively. Interestingly, we observed that these different populations had unique haplotypes varying with their habitats, demonstrating sequence divergence in the homogenous population of A. franciscana despite them being in a similar geographic zone.
... The mitochondrial markers, COX1 and 16S, have been successfully used in phylogeny of branchiopods [60][61][62][63][64][65]. To date only COX1 has been utilized for phylogenetic studies on Artemia, nevertheless mitogenomic results demonstrated significant difference in the nucleotide composition of ATP8, ATP6, ND3, ND6, ND1 and COX3. ...
Article
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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). ...
... Our results also confirmed the colonization of the same species in GL. SFB and GSL are the two major sources of Artemia that are usually used to culture in saline ecosystems for industrial aquaculture and fishery activities to produce Artemia cysts and biomass (Eimanifar et al., 2014;Muñoz et al., 2014;Asem et al., 2018;Saji et al., 2019). Thus, these populations were considered in this analysis to determine the genetic alterations of colonized populations in new nonnative environments. ...
... Mitochondrial DNA presents some exceptional characteristics, including rapid evolutionary rates, maternal origin, and lack of recombination (Boore, 1999;Miller et al., 2009). Therefore, mitochondrial markers are important for the apprehension of tracing and the explanation of the source of non-indigenous species in new habitats (Ashton et al., 2008;Ficetola et al., 2008;Mabuchi et al., 2008;Gaubert et al., 2009;Asem et al., 2018;Saji et al., 2019). ...
Article
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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.