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Maximum parsimony haplotype network of ITS1 sequences. The size of each square/circle is proportional to the frequency of specimens. Each joining line between haplotypes is equal with single nucleotide substitutions. Black dots between haplotypes. Full-size DOI: 10.7717/peerj.7190/fig-4
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Urmia Lake, the largest natural habitat of the brine shrimp Artemia urmiana, has progressively desiccated over the last two decades, resulting in a loss of 80% of its surface area and producing thousands of hectares of arid salty land. This ecological crisis has seriously affected the lake's native biodiversity. Artemia urmiana has lost more than 9...
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Context 1
... for 58 specimens revealed 56 haplotypes (H1-H56), which showed high variation in comparison with the other markers. The central haplotype (H1) covered only two specimens (3.5%) including a single specimen from each year. With the exception of H39 that was shared by two specimens from 2004, other haplotypes were unique to a single specimen (Fig. ...
Context 2
... 16S structure. Tajima's D and Fu and Li's D* were non-significant in the rainy period which could indicate 16S marker was at demographic equilibrium without selection in the rainy period. In contrast, a negative and significant neutrality value and expanded haplotype network indicated that 16S is involved in recent expansion in the drought period (Fig. 6). The negative and significant value of Fu Fs test suggested the excess of new mutations in the gene pool of 16S in both normal and drought ...
Citations
... Based on COI and ISSR markers, Eimanifar and Wink (2013) documented that the Urmia Lake population exhibited a high level of haplotype diversity, a low level of nucleotide diversity, and no apparent genetic structure. Asem et al. (2019) found that the increased salinity of Urmia Lake during 1994-2004 had caused changes in the genetic diversity and genetic structure of this Artemia population. Although there are many Artemia populations in the QTP, population genetic analysis was only documented for single population (the Shuanghu population; Han et al., 2019). ...
... Genomic DNA was extracted from single resting egg using Chelex® 100 resin (Bio-Rad Laboratories, Hercules, CA, USA) as described by Asem et al. (2019). ...
Bird-mediated dispersal of resting eggs is the main mechanism for Artemia dispersal among catchments. The
bisexual populations of Artemia urmiana species complex, which is here considered to be a collection of Artemia
genetically close to the so-called “Western Asian Lineage”, are mostly distributed in central and western Asia (i.
e., in regions falling into the Central Asian Flyway of migratory birds) and live in diversified habitats. Little is
known about the genetic relationships among these populations. Aiming to understand the population genetic
characteristics and the roles of migratory birds on the dispersal and gene flow of this Artemia group, we evaluated
the genetic diversity, genetic differentiation, and gene flow among 14 populations, with their altitudes ranging
from 540 to 4870 m above sea level, using 13 microsatellite markers. Almost all populations exhibited high
genetic diversity and heterozygote excess, which may be a consequence of combined effects of dispersal and
hybridization. The global genetic differentiation (FST) value was 0.092, the pairwise FST values were
0.003–0.246. Discriminant analysis of principal components identified three genetic clusters, consisting of Urmia
Lake (Iran), Zhundong (Xinjiang, China), and 12 Qinghai-Tibet Plateau populations, respectively. The amongpopulation
genetic differentiation seems to be a consequence of isolation by distance and adaptation to diversified
habitats induced by altitudinal gradient. Historical gene flows are asymmetrical, and show an evolutionary
source-sink dynamics, with Jingyu Lake (Xinjiang, China) population being the major source. These results
support our hypothesis that in Qinghai-Tibet Plateau and surrounding areas the bird-mediated dispersal of
Artemia may be biased towards from north to south and/or from higher altitude to lower altitude.
... Located in northwest Iran, Urmia Lake is one of the international wetlands registered by the Ramsar Convention. It is home to the largest natural habitat of Artemia species of brine shrimp [39] and a location with associated ecotourism potential [40]. The wetland has shrunk and lost much of its area over the past decade. ...
This review paper investigates unprecedented recent investment and capital spending in cross-provincial water management and transfer infrastructure in Iran. Although numerous cross-provincial water transfer plans have been implemented in the country, the scale, approach, and stakeholders involved in this recent national plan are unprecedented. This notable national plan includes long water transfer pipelines that pass through seven provinces, aiming at transferring desalinated water from the Persian Gulf to the interior. Regarding the scale, there have been a few cases of transferring water at the cross-provincial level, but mostly across two provinces. The approach has also changed in this plan. Long-term efforts to construct dams for electricity or farming with limited geographical impact have been shifted to desalinating water, long-distance pipe-laying, large-scale electric-powered water pumping, and long-distance underground tunnels. Stakeholders have become more diverse, from the council members of a small village to high-level decision makers at the national level. This paper elaborates on these extraordinary alterations by exemplifying a few other contemporary case studies of water transfer plans in Iran. It also examines the fundamental logic, bottlenecks, and future scenarios of this large-scale plan.
... Compared with marine and freshwater environments, the diversity and number of species in high-salinity habitat systems are often limited, which makes it more vulnerable and poses greater threats to biodiversity within this ecosystems. Since salinity is the main environmental factor for the survival and population reproduction of Artemia [9], a study found that the gradual drying of Lake Urmia in Iran has led to near-saturation levels of lake salinity and the loss of more than 90% of Artemia populations over the past two decades [11]. Similarly, the Aral Sea, the largest saltlake in central Asia, is also facing shrinking water levels and rising salinity, leading to a decline in Artemia cysts production. ...
Brine shrimp Artemia is the crucial live food in fish and crustacean larviculture. With the decrease of Artemia resources and the increase in aquaculture demand, it is necessary to conduct Artemia population genetics and to manage this important resource efficiently. In this study, the genetic diversity and distribution pattern of Artemia populations originated from ten inland salt lakes in central-west China were studied using a large number of single nucleotide polymorphisms (SNPs) obtained by high-throughput sequencing. The results showed that Tibetan populations had the highest level of genetic diversity, while Shanxi population had the lowest value. Genetic structure and principal component analysis further revealed these populations can be classified into four geographically and genetically distinct groups, and Tibetan populations were further divided into two subgroups and showed a trend of higher in the west and lower in the east at the level of genetic diversity. In addition, a certain amount of gene flow was observed among these 10 populations, and A. sinica had a unidirectional gene flow to all populations in the eastern Nagri region. Finally, species distribution models indicated that Bio1, Bio4, and Bio12 were the main environmental factors affecting the distribution of Artemia , and climate fluctuations had a strong influence on the distribution of Artemia . The findings revealed Tibetan populations will be further reduced in the future, and it is necessary to adopt a series of measures to strengthen Artemia protection and promote rational exploitation and utilization of this natural resources. This study further improves our understanding of the genetic structure of Artemia in central-west China, and provides certain guidance for the protection of Artemia germplasm resources.
... Several studies demonstrate the effect of ecological conditions on aquatic organism genetic variation (Paul et al. 2012, Stoks et al. 2014, Asem et al. 2019, Bernal et al. 2022. Previous studies show that A. urmiana has widely distributed haplotype networks and great genetic variation , 2020, Asem et al. 2019. ...
... Several studies demonstrate the effect of ecological conditions on aquatic organism genetic variation (Paul et al. 2012, Stoks et al. 2014, Asem et al. 2019, Bernal et al. 2022. Previous studies show that A. urmiana has widely distributed haplotype networks and great genetic variation , 2020, Asem et al. 2019. High genetic variation in A. urmiana is attributed to extensive historical fluctuations in ecological conditions with impacts on effective population sizes in Urmia Lake (Asem et al. 2012(Asem et al. , 2019 driving the fixation of new mutations (Asem et al. 2019). ...
... Previous studies show that A. urmiana has widely distributed haplotype networks and great genetic variation , 2020, Asem et al. 2019. High genetic variation in A. urmiana is attributed to extensive historical fluctuations in ecological conditions with impacts on effective population sizes in Urmia Lake (Asem et al. 2012(Asem et al. , 2019 driving the fixation of new mutations (Asem et al. 2019). Similar results are known from other anostracan families as well (e.g. ...
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.
... Since Lake Urmia is a closed basin, the main causes of this pronounced 8-m drop in lake level were likely a decrease in riverine inflow and/or excess evaporative outflow, which could have been caused by anthropogenic alterations including dam construction, channel deviation, groundwater pumping (Tourian et al., 2015), as well as climate change involving reduced precipitation and/or increased temperatures. Although anthropogenic factors have been widely discussed (Ahmady-Birgani et al., 2020;Asem et al., 2019;Chaudhari et al., 2018;Khazaei et al., 2019), the lack of high-resolution sedimentary records in the past limits our understanding of ongoing processes in the context of present-day climate variations. Existing data either cover a very long time scale but with low resolution, or are discontinuous, or concern a very short-term period (Djamali et al., 2008a;Kelts and Shahrabi, 1986;Stevens et al., 2012;Talebi et al., 2016). ...
... Among them, temporary ecosystems are especially vulnerable (Zacharias and Zamparas 2010). Climatic change and the resulting unpredictable rainfall distribution threaten the fill of endorheic basins, halting the formation of the necessary aquatic habitats for development of anostracans (Asem et al. 2019;Rautio et al. 2011) and other invertebrates (Williams 1997). Similarly, changes in light and temperature alter hatching success in Anostraca (Tladi et al. 2020). ...
The fairy shrimp Branchinectella media, because of its passive dispersal capacity and scarce and irregularly distributed habitats (temporary saline aquatic systems), is an intriguing organism from a population genomics and conservation per- spective. Stochasticity of dispersal events and the irregular distribution of its habitat might lead to low levels of population connectivity and genetic diversity, and consequently, populations with limited persistence through time. Indeed, by using genomic data (SNPs), we found a strong genetic structure among some of the geographically isolated Iberian populations of B. media. Interestingly, we also obtained high estimates of effective population sizes. Lack of suitable habitat between populations (absence of a “stepping stone” network) and strong genetic differentiation suggest limited dispersal success in B. media. However, the high effective population sizes observed ensure persistence of B. media populations against genetic stochasticity (genetic drift). These results indicate that rescue-effect might not be essential for population persistence if they maintain high effective population sizes able to hold adequate levels of genetic diversity. Should high population sizes be reported in other low dispersing Anostraca, one might be optimistic with regard to their conservation status and fate, provided that their natural habitats remain undisturbed.
... Regarding genetic diversity in COI, A. urmiana exhibited high variation in agreement with other studies (Eimanifar et al., 2014, and the highest interspecific variation among all the markers considered in this study, whereas A. tibetiana and A. amati n. sp. had the lowest variation. Negative and significant values of the D Tajima test in A. urmiana and A. tibetiana considering mitochondrial markers (except 12S) suggest the existence of rare alleles at high frequencies, coincidently with a population expansion after a probable bottleneck or the effect of selection (Asem et al., 2019b). ...
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.
... 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). ...
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.
... In the early 1990s, Artemia (Artemia parthenogenetica) was discovered in the Aral Sea, and since the 2000s it has become a permanent and dominant component of the reservoir, accounting for more than 99% of the total zooplankton biomass [5]. However, many questions of the biology of the emerging population of this reservoir in Central Asia have not been studied yet. ...
Artemia (Artemia parthenogenetica) has been consistently identified in the plankton of the salinizing Aral Sea since the 1990s and by now it has become dominant in the water body. The objective of the research was to investigate diameter, population and distribution of Artemia cysts in Aral Sea Basin, Uzbekistan. During the study observation, the diameter of untreated and decapsulated cysts and the chorion thickness were determined in samples collected in November 2020 on the eastern and southern shores of the Aral Sea remnant (Uzbekistan). The results showed that in samples from the eastern coast, the sizes of untreated cysts were 210-310 (on average 270.6 + 3.1) µm, the sizes of decapsulated cysts were 160-260 (212.2 + 3.8) µm, the calculated diameter of the chorion was 29.2 µm. In samples from the southern coast, the sizes of untreated cysts were 198-330 (265.3 + 4.09) µm, the sizes of decapsulated cysts were 198-297 (242.8 + 3.37) µm, and the calculated diameter of the chorion was 11.1 µm.
... Our results revealed a negative and significant Tajima's D value (−2.304) only for the exotic population in AWWR (Table 6), which demonstrated an excess of rare haplotypes followed by population expansion or purifying selection (Nei and Kumar, 2000;Swanson et al., 2001;Akey et al., 2004;Cruciani et al., 2008;Levitan and Stapper, 2009;Asem et al., 2019). Given the greater numbers of polymorphic sites (12 sites), mutations (12 mutations), and haplotypes (12 haplotypes) in comparison with other localities, Tajima's D result might be ascribed to the demographic expansion of the Artemia population in AWWR. ...
... The negative values of Fu's Fs test and Fu and Li's D* test suggest the existence of rare recent mutations (Fu, 1997;Ramos-Onsins and Rozas, 2002;Zhao et al., 2008;Asem et al., 2019) and an excess of rare historical mutations in populations (Fu and Li, 1993;Fu, 1996;Zhao et al., 2008;Asem et al., 2019), respectively. The results of both neutrality tests were negative and significant for the AWWR population. ...
... The negative values of Fu's Fs test and Fu and Li's D* test suggest the existence of rare recent mutations (Fu, 1997;Ramos-Onsins and Rozas, 2002;Zhao et al., 2008;Asem et al., 2019) and an excess of rare historical mutations in populations (Fu and Li, 1993;Fu, 1996;Zhao et al., 2008;Asem et al., 2019), respectively. The results of both neutrality tests were negative and significant for the AWWR population. ...
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.
