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Names that have ever been used for native brine shrimps Artemia of Asia, their availability, infor- mation of type specimens and nomenclatural problems

Names that have ever been used for native brine shrimps Artemia of Asia, their availability, infor- mation of type specimens and nomenclatural problems

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The genus Artemia Leach, 1819 is a cosmopolitan halophilic crustacean, consisting of bisexual species and obligate parthenogenetic populations. Asia is rich in Artemia biodiversity. More than 530 Artemia sites have been recorded from this area and more than 20 species/subspecies/variety names have been used for them. There exist various problems in...

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Context 1
... far as we are aware, 14 binomens (including the one questionably reported as a species of Branchinecta Verrill, 1869) and nine trinomens, as well as unidentified species/subspecies/varieties, have appeared in the form of scientific names (combined with a genus name and typeset in italics) in literature relating to native Asian brine shrimps. As shown in Table 1, almost all of the names have some kind of nomenclatural problem. Among the 13 binomens proposed for Artemia, eleven fulfil the availability requirements of International Code of Zoological Nomenclature (ICZN, 4 th edition) and are considered to be available species names [Artemia salina (Lin- naeus, 1758); Artemia asiatica Walter, 1887;Artemia urmiana Günther, 1899;Ar- temia parthenogenetica Bowen & Sterling, 1978;Artemia sinica Cai, 1989;Artemia barkolica Qian and Wang in Qian et al. 1992; Artemia urumuqinica Qian and Wang in Qian et al. 1992; Artemia ebinurica Qian and Wang in Qian et al. 1992; Artemia tibetiana Abatzopoulos, Zhang & Sorgeloos, 1998;Artemia frameshifta Naganawa & Mura, 2017; Artemia murae Naganawa in Naganawa and Mura 2017], and the other two are unavailable [Artemia kazakhastan Vikas et al., 2012;Artemia china Vikas et al., 2012]. ...
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... two decades, Tibetan bisexual populations have been considered as belonging to a single species, A. tibetiana, originally described as a bisexual species from Lagkor Co, Tibet, China ( Abatzopoulos et al. 1998). However, as that in Urmia Lake, a parthenogenetic population was also documented from this lake (Van Stappen et al. 2003;Maccari et al. 2013; see Table 1). Wang et al. (2008) documented four Tibetan bisexual populations clustering in two different clades using the mitochondrial COI marker, with one clade only hosting the type locality population (Lagkor Co) and a second distinct clade hosting the others (Kyêbxang Co (=Qixiang Lake or Qi Xiang Cuo), Nima, and Yangnapeng Co). ...
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... other bisexual populations, the Jingyu Lake (Xinjiang, China) population and the Xiao Qaidam Lake (Qinghai, China) population, were considered to represent different subspecies of A. sinica. The subspecies names Artemia sinica jingyuhuensis Yin, Zhang & You, 2013 andArtemia sinica xiaochaidanensis Yin, Zhang & You, 2013 were proposed for them, respectively, though they are not available (Table 1). Zheng and Sun (2008) documented some morphological differences between Jingyu Lake and Lagkor Co populations though they identified the former population as A. tibetiana. ...
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... nomenclatural and taxonomic status of Artemia parthenogenetica have been discussed by previous authors (Barigozzi 1980;Belk and Brtek 1995;Baxevanis et al. 2006; see also Table 1). Since Bowen and Sterling (1978), this name has been used in numerous publications (sometimes followed by a population site), whereas some authors preferred to refer parthenogenetic Artemia as "populations" ( Abatzopoulos et al. 2002a;Baxevanis et al. 2006;Asem et al. 2010). ...
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... far as we are aware, 14 binomens (including the one questionably reported as a species of Branchinecta Verrill, 1869) and nine trinomens, as well as unidentified species/subspecies/varieties, have appeared in the form of scientific names (combined with a genus name and typeset in italics) in literature relating to native Asian brine shrimps. As shown in Table 1, almost all of the names have some kind of nomenclatural problem. Among the 13 binomens proposed for Artemia, eleven fulfil the availability requirements of International Code of Zoological Nomenclature (ICZN, 4 th edition) and are considered to be available species names [Artemia salina (Lin- naeus, 1758); Artemia asiatica Walter, 1887;Artemia urmiana Günther, 1899;Ar- temia parthenogenetica Bowen & Sterling, 1978;Artemia sinica Cai, 1989;Artemia barkolica Qian and Wang in Qian et al. 1992; Artemia urumuqinica Qian and Wang in Qian et al. 1992; Artemia ebinurica Qian and Wang in Qian et al. 1992; Artemia tibetiana Abatzopoulos, Zhang & Sorgeloos, 1998;Artemia frameshifta Naganawa & Mura, 2017; Artemia murae Naganawa in Naganawa and Mura 2017], and the other two are unavailable [Artemia kazakhastan Vikas et al., 2012;Artemia china Vikas et al., 2012]. ...
Context 6
... two decades, Tibetan bisexual populations have been considered as belonging to a single species, A. tibetiana, originally described as a bisexual species from Lagkor Co, Tibet, China ( Abatzopoulos et al. 1998). However, as that in Urmia Lake, a parthenogenetic population was also documented from this lake (Van Stappen et al. 2003;Maccari et al. 2013; see Table 1). Wang et al. (2008) documented four Tibetan bisexual populations clustering in two different clades using the mitochondrial COI marker, with one clade only hosting the type locality population (Lagkor Co) and a second distinct clade hosting the others (Kyêbxang Co (=Qixiang Lake or Qi Xiang Cuo), Nima, and Yangnapeng Co). ...
Context 7
... other bisexual populations, the Jingyu Lake (Xinjiang, China) population and the Xiao Qaidam Lake (Qinghai, China) population, were considered to represent different subspecies of A. sinica. The subspecies names Artemia sinica jingyuhuensis Yin, Zhang & You, 2013 andArtemia sinica xiaochaidanensis Yin, Zhang & You, 2013 were proposed for them, respectively, though they are not available (Table 1). Zheng and Sun (2008) documented some morphological differences between Jingyu Lake and Lagkor Co populations though they identified the former population as A. tibetiana. ...
Context 8
... nomenclatural and taxonomic status of Artemia parthenogenetica have been discussed by previous authors (Barigozzi 1980;Belk and Brtek 1995;Baxevanis et al. 2006; see also Table 1). Since Bowen and Sterling (1978), this name has been used in numerous publications (sometimes followed by a population site), whereas some authors preferred to refer parthenogenetic Artemia as "populations" ( Abatzopoulos et al. 2002a;Baxevanis et al. 2006;Asem et al. 2010). ...

Citations

... 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]. ...
... However, results of morphological and genetic investigations were contradictory [22]. To date, the taxonomy and biosystematics of Artemia are still controversial, especially concerning the Asian species [21]. Mitogenomic information could provide a better reconstruction of the maternal evolutionary mechanism and phylogenetic status of Artemia. ...
... Generally, some of the adjacent lagoons are connected with Urmia Lake when the lake level raises annually during rainy seasons in spring and autumn [29], which increases the probability of collecting parthenogenetic specimens along the shoreline of the lake (Atashbar, personal communication, Urmia University). On the other hand, genetic variation between parthenogenetic populations and A. urmiana is quite low [19,21]. Additionally, a current study based on barcoding with the mitochondrial COX1 marker found that parthenogenetic populations in some localities share same haplotypes with A. urmiana [30]. ...
Article
Full-text available
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.
... The brine shrimp Artemia is a member of the branchiopod crustaceans and is distributed worldwide in various saline habitats [29]. In the genus, seven species undergo bisexual reproduction, and numerous parthenogenetic populations, also known as A. parthenogenetica, can reproduce asexually [30][31][32]. Female and male Artemia are easily distinguished by their phenotypes. Females are usually 1-2 mm larger than males and have an ovisac. ...
Article
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The brine shrimp Artemia has a ZW sex determination system with ZW chromosomes in females and ZZ chromosomes in males. Artemia has been considered a promising model organism for ZW sex-determining systems, but the genes involved in sex determination and differentiation of Artemia have not yet been identified. Here, we conducted transcriptome sequencing of female and male A. franciscana using PacBio Iso-Seq and Illumina RNA-Seq techniques to identify candidate sex determination genes. Among the 42,566 transcripts obtained from Iso-Seq, 23,514 were analyzed. Of these, 2065 (8.8%) were female specific, 2513 (10.7%) were male specific, and 18,936 (80.5%) were co-expressed in females and males. Based on GO enrichment analysis and expression values, we found 10 female-biased and 29 male-biased expressed genes, including DMRT1 and Sad genes showing male-biased expression. Our results showed that DMRT1 has three isoforms with five exons, while Sad has seven isoforms with 2–11 exons. The Sad gene is involved in ecdysteroid signaling related to molting and metamorphosis in arthropods. Further studies on ecdysteroid biosynthetic genes are needed to improve our understanding of Artemia sex determination. This study will provide a valuable resource for sex determination and differentiation studies on Artemia and other crustaceans with ZW systems.
... Artemia is a poorly diversified group of small hypersaline water branchiopods (Crustacea, Anostraca), currently conformed by less than a dozen species distributed all over the world, often associated to salt production, and used as a model system for diverse research purposes, as well as a valuable food source in aquaculture (Lenz, 1984;Sorgeloos et al., 1986;Sorgeloos, Dhert & Candreva, 2001;Van Stappen, 1996;Ruebhart, Cock & Shaw, 2008;Amat et al., 2005;Baxevanis, Kappas & Abatzopoulos, 2006). Despite the reduced number of species, the different taxa within Artemia have been referred to, in the scientific literature, with more than 50 names, almost all of them used at the species level (Daday de Deés, 1910;Belk & Brtek, 1995;Rogers, 2013;Asem et al., 2020). Most of the names applied from the end of the eighteen to the mid-twentieth century in Artemia taxonomic characterization were forgotten and not used again by later authors. ...
... It is difficult to believe that a proper revision of the nomenclature in accordance to the rules and recommendations of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1999) has not been performed yet for one of the most world-wide commercialized invertebrates. Only Asem, Rastegar-Pouyani & De Los Ríos-Escalante (2010) made a clarification attempt, and recently, Asem et al. (2020) reviewed the taxonomic problems of native Asian Artemia. The task has been probably avoided either because the early inclusion of partial genetic data in the definition of taxa blurred the overall picture (Alonso, 1996), or because the early proliferation of names made the selection of valid names for the molecularly defined taxa a complicated task. ...
... An exhaustive bibliographical search was undertaken to locate and gather all original publications in which any possible nomenclatural act affecting Artemia was published. The search started with four main sources for synonymies: Daday de Deés (1910), Belk & Brtek (1995), Asem, Rastegar-Pouyani & De Los Ríos-Escalante (2010), Asem et al. (2020), andRogers (2013). From there, we sought for any additional bibliographic information mentioned in each of the papers consulted. ...
Article
Full-text available
High morphological plasticity in populations of brine shrimp subjected to different environmental conditions, mainly salinity, hindered for centuries the identification of the taxonomic entities encompassed within Artemia . In addition, the mismatch between molecular and morphological evolution rates complicates the characterization of evolutionary lineages, generating taxonomic problems. Here, we propose a phylogenetic hypothesis for Artemia based on two new complete mitogenomes, and determine levels of congruence in the definition of evolutionary units using nuclear and mtDNA data. We used a fossil of Artemia to calibrate the molecular clock and discuss divergence times within the genus. The hypothesis proposed herein suggests a more recent time frame for lineage splitting than previously considered. Phylogeographic analyses were performed using GenBank available mitochondrial and nuclear markers. Evidence of gen e flow, identified through discordances between nuclear and mtDNA markers, was used to reconsider the specific status of some taxa. As a result, we consider Artemia to be represented by five evolutionary units: Southern Cone, Mediterranean—South African, New World, Western Asian, and Eastern Asian Lineages. After an exhaustive bibliographical revision, unavailable names for nomenclatural purposes were discarded. The remaining available names have been assigned to their respective evolutionary lineage. The proper names for the evolutionary units in which brine shrimps are structured remain as follows: Artemia persimilis Piccinelli & Prosdocimi, 1968 for the Southern Cone Lineage, Artemia salina (Linnaeus, 1758) for the Mediterranean-SouthAfrican Lineage, Artemia urmiana Günther, 1899 for the Western Asian Lineage, and Artemia sinica Cai, 1989 for the Eastern Asian Lineage. The name Artemia monica Verrill, 1869 has nomenclatural priority over A. franciscana Kellogg, 1906 for naming the New World Lineage. New synonymies are proposed for A. salina ( = C. dybowskii Grochowski, 1896 n. syn. , and A. tunisiana Bowen & Sterling, 1978 n. syn. ), A. monica (= A. franciscana Kellogg, 1906 n. syn ., and A. salina var. pacifica Sars, 1904 n. syn. ); A. urmiana (= B. milhausenii Fischer de Waldheim, 1834 n. syn. , A. koeppeniana Fischer, 1851 n. syn. , A. proxima King, 1855 n. syn. , A. s. var. biloba Entz, 1886 n. syn. , A. s. var. furcata Entz, 1886 n. syn. , A. asiatica Walter, 1887 n. syn. , A. parthenogenetica Bowen & Sterling, 1978 n. syn. , A. ebinurica Qian & Wang, 1992 n. syn. , A. murae Naganawa, 2017 n. syn. , and A. frameshifta Naganawa & Mura, 2017 n. syn. ). Internal deep nuclear structuring within the A. monica and A. salina clades, might suggest the existence of additional evolutionary units within these taxa.
... "parthenogenetic strain" ( Table 1). The taxonomy and nomenclature of Artemia parthenogenetic populations is still unsettled (e.g., Asem et al. 2020;Sainz-Escudero et al. 2021), and the binomen Artemia parthenogenetica itself is considered a nomen dubium by Rogers (2013). ...
... 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]. ...
... However, results of morphological and genetic investigations were contradictory [22]. To date, the taxonomy and biosystematics of Artemia are still controversial, especially concerning the Asian species [21]. Mitogenomic information could provide a better reconstruction of the maternal evolutionary mechanism and phylogenetic status of Artemia. ...
... Generally, some of the adjacent lagoons are connected with Urmia Lake when the lake level raises annually during rainy seasons in spring and autumn [29], which increases the probability of collecting parthenogenetic specimens along the shoreline of the lake (Atashbar, personal communication, Urmia University). On the other hand, genetic variation between parthenogenetic populations and A. urmiana is quite low [19,21]. Additionally, a current study based on barcoding with the mitochondrial COX1 marker found that parthenogenetic populations in some localities share same haplotypes with A. urmiana [30]. ...
Article
Full-text available
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.
... The genus Artemia consists of seven bisexual species and a large number of parthenogenetic populations with different ploidy levels [3,11]. It has been assumed that Asia is the origin of parthenogenetic populations [3,12]. Recently, two new species, Artemia frameshifta and Artemia murae have been described from Mongolia [13]. ...
Article
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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 (ISSRs) to determine the genetic variability and population structure of Artemia populations (indigenous and introduced) from 14 different geographical locations in Western Asia. Based on the haplotype spanning network, Artemia urmiana has exhibited higher genetic variation than native parthenogenetic populations. Although A. urmiana represented a completely private haplotype distribution, no apparent genetic structure was recognized among the native parthenogenetic and invasive A. franciscana populations. Our ISSR findings have documented that despite that invasive populations have lower variation than the source population in Great Salt Lake (Utah, USA), they have significantly revealed higher genetic variability compared to the native populations in Western Asia. According to the ISSR results, the native populations were not fully differentiated by the PCoA analysis, but the exotic A. franciscana populations were geographically divided into four genetic groups. We believe that during the colonization, invasive populations have experienced substantial genetic divergences, under new ecological conditions in the non-indigenous regions.
Article
We document morphology and genetic relatedness of morphotypes within the Artemia franciscana that has colonized Covelong saltern in Kelambakkam (South India). They exhibits high phenotypic plasticity and intra-population variation in a supposedly panmictic population. Morphometric analysis was carried out on 18 traits in males and 15 in females and subjected to multivariate analysis. Three types each in males (M1, M3 and M4) and females (F1, F2 and F3) were singled out. The axes in the plot explain 77.9% in males and 73.9% in females of the variation through discriminant function analysis. The sequence divergence of the population was evaluated through mtDNA (16S rRNA) and nDNA (p26 gene) markers. The substantial sequence divergence was observed between the morphotypes through nDNA and appearing sequence divergence was recognized by mtDNA. Likewise, phylogenetic analyses of maximum likelihood and Bayesian analysis through nDNA revealed a maximum similarities between M1, M4 with F1 & F2 and M3 with F3. Further, haplotype distribution of nDNA revealed unique haplotype between M1 and M4 with F1 and F2 morphotypes. This study thus confirms the occurrence of morphotypes with specific characteristics inside a seemingly homogeneous population.
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Due to the lack of a taxonomic key for the identification of Artemia species, molecular markers have been increasingly used for phylogenetic studies. The mtCOI marker is a regularly considered marker for the molecular systematics of Artemia populations. The proposed universal and specific primers have mostly failed to amplify the Artemia aff. sinica mtCOI marker, and on the whole, the successfully amplified products of the PCR were inefficient, primarily through the representation of poly-peak or incomplete sequences. We presumed that if a forward primer could be developed regarding the joint regions of the last part of the previous gene (tRNATyr) and the beginning of the target gene mtCOI, the sequence could be relevant to the target-sequence of mtCOI. Thus, here, we describe a new set of primers, which could be used to amplify the mtCOI barcoding region of Artemia aff. sinica Cai, 1989, with a high performance of sequencing. The new primer set worked well also for other Artemia bisexual species, as well as for parthenogenetic populations. It is recommended that joint regions between the previous/next gene(s) and the target marker, could be aimed at when designing specific primers for other markers and taxa.
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Determining how and how often asexual lineages emerge within sexual species is central to our understanding of sex-asex transitions and the long-term maintenance of sex. Asexuality can arise "by transmission" from an existing asexual lineage to a new one, through different types of crosses. The occurrence of these crosses, cryptic sex, variation in ploidy and recombination within asexuals greatly complicates the study of sex-asex transitions, as they preclude the use of standard phylogenetic methods and genetic distance metrics. In this study we show how to overcome these challenges by developing new approaches to investigate the origin of the various asexual lineages of the brine shrimp Artemia parthenogenetica. We use a large sample of asexuals, including all known polyploids, and their sexual relatives. We combine flow cytometry with mitochondrial and nuclear DNA data. We develop new genetic distance measures and methods to compare various scenarios describing the origin of the different lineages. We find that all diploid and polyploid A. parthenogenetica likely arose within the last 80,000 years through successive and nested hybridization events that involved backcrosses with different sexual species. All A. parthenogenetica have the same common ancestor and therefore likely carry the same asexuality gene(s) and reproduce by automixis. These findings radically change our view of sex-asex transitions in this group, and show the importance of considering asexuality "by transmission" scenarios. The methods developed are applicable to many other asexual taxa.