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1.  A revision of Artemia biodiversity in Macaronesia 
Aquatic Biosystems  2012;8:25.
In a biogeographical context, the term Macaronesia broadly embraces the North Atlantic archipelagos of the Azores, Madeira, Selvagens, the Canary Islands, and Cape Verde. The peculiar arid climatic conditions in some of these places have led to the development of marine salt exploitations, which can be counted among the hypersaline habitats of the brine shrimp Artemia (Branchiopoda, Anostraca). Parthenogenetic populations of this anostracan were described in the Canary Islands during the last decades of the 20th century, while the American Artemia franciscana species was recently found in the Cape Verde archipelago. Following an invasive pattern, this exotic species has recently reached the Canary Islands, too. This paper reports information dealing with biotope loss (solar saltworks) in this biogeographical region, together with possible consequences concerning the arrival of invasive species, two factors that frequently promote dramatic biodiversity losses. The discussion of this threat focuses mainly on the Canary Islands archipelago where native species of Artemia still exist.
PMCID: PMC3543279  PMID: 23075404
Artemia; Biodiversity; Macaronesia; Saltworks; Loss of habitats; Invasion
2.  The Brine Shrimp Artemia: Adapted to Critical Life Conditions 
The brine shrimp Artemia is a micro-crustacean, well adapted to the harsh conditions that severely hypersaline environments impose on survival and reproduction. Adaptation to these conditions has taken place at different functional levels or domains, from the individual (molecular-cellular-physiological) to the population level. Such conditions are experienced by very few equivalent macro-planktonic organisms; thus, Artemia can be considered a model animal extremophile offering a unique suite of adaptations that are the focus of this review. The most obvious is a highly efficient osmoregulation system to withstand up to 10 times the salt concentration of ordinary seawater. Under extremely critical environmental conditions, for example when seasonal lakes dry-out, Artemia takes refuge by producing a highly resistant encysted gastrula embryo (cyst) capable of severe dehydration enabling an escape from population extinction. Cysts can be viewed as gene banks that store a genetic memory of historical population conditions. Their occurrence is due to the evolved ability of females to “perceive” forthcoming unstable environmental conditions expressed by their ability to switch reproductive mode, producing either cysts (oviparity) when environmental conditions become deleterious or free-swimming nauplii (ovoviviparity) that are able to maintain the population under suitable conditions. At the population level the trend is for conspecific populations to be fragmented into locally adapted populations, whereas species are restricted to salty lakes in particular regions (regional endemism). The Artemia model depicts adaptation as a complex response to critical life conditions, integrating and refining past and present experiences at all levels of organization. Although we consider an invertebrate restricted to a unique environment, the processes to be discussed are of general biological interest. Finally, we highlight the benefits of understanding the stress response of Artemia for the well-being of human populations.
PMCID: PMC3381296  PMID: 22737126
Artemia; animal extremophile; evolution; adaptation; salty lakes; stressful environments
3.  Variable hydrology and salinity of salt ponds in the British Virgin Islands 
Saline Systems  2006;2:2.
Caribbean salt ponds are unique wetlands that have received little scientific attention. They are common features of dry Caribbean coastlines, but they are threatened by rapid coastal development. We compared hydrology and salinity of 17 salt ponds in the British Virgin Islands. Ponds were mostly hypersaline (>50 ppt), and they exhibited dramatic salinity fluctuations in response to rainfall and evaporation. Individual ponds varied in their mean salinities and thus experienced different ranges of salinity. Differences in mean salinity appeared to be linked with hydrological characteristics. Hydrological variation ranged from permanently inundated ponds with direct sea connection to those fully isolated from the sea and retaining water only after rainfall. We characterized groups of ponds by their major hydrological characteristics, particularly their period of inundation and their degree of connection with the sea. The resulting classification appeared to reflect a continuum of increasing isolation from the sea, concurring with published geological records from salt pond sediments elsewhere. The patterns of variability and succession described here are applicable to salt pond management interests throughout the Caribbean.
PMCID: PMC1413541  PMID: 16504156
4.  Saline systems of the Great Plains of western Canada: an overview of the limnogeology and paleolimnology 
Saline Systems  2005;1:10.
In much of the northern Great Plains, saline and hypersaline lacustrine brines are the only surface waters present. As a group, the lakes of this region are unique: there is no other area in the world that can match the concentration and diversity of saline lake environments exhibited in the prairie region of Canada and northern United States. The immense number of individual salt lakes and saline wetlands in this region of North America is staggering. Estimates vary from about one million to greater than 10 million, with densities in some areas being as high as 120 lakes/km2.
Despite over a century of scientific investigation of these salt lakes, we have only in the last twenty years advanced far enough to appreciate the wide spectrum of lake types, water chemistries, and limnological processes that are operating in the modern settings. Hydrochemical data are available for about 800 of the lake brines in the region. Composition, textural, and geochemical information on the modern bottom sediments has been collected for just over 150 of these lakes. Characterization of the biological and ecological features of these lakes is based on even fewer investigations, and the stratigraphic records of only twenty basins have been examined.
The lake waters show a considerable range in ionic composition and concentration. Early investigators, concentrating on the most saline brines, emphasized a strong predominance of Na+ and SO4-2 in the lakes. It is now realized, however, that not only is there a complete spectrum of salinities from less than 1 ppt TDS to nearly 400 ppt, but also virtually every water chemistry type is represented in lakes of the region. With such a vast array of compositions, it is difficult to generalize. Nonetheless, the paucity of Cl-rich lakes makes the northern Great Plains basins somewhat unusual compared with salt lakes in many other areas of the world (e.g., Australia, western United States). Compilations of the lake water chemistries show distinct spatial trends and regional variations controlled by groundwater input, climate, and geomorphology. Short-term temporal variations in the brine composition, which can have significant effects on the composition of the modern sediments, have also been well documented in several individual basins.
From a sedimentological and mineralogical perspective, the wide range of water chemistries exhibited by the lakes leads to an unusually large diversity of modern sediment composition. Over 40 species of endogenic precipitates and authigenic minerals have been identified in the lacustrine sediments. The most common non-detrital components of the modern sediments include: calcium and calcium-magnesium carbonates (magnesian calcite, aragonite, dolomite), and sodium, magnesium, and sodium-magnesium sulfates (mirabilite, thenardite, bloedite, epsomite). Many of the basins whose brines have very high Mg/Ca ratios also have hydromagnesite, magnesite, and nesquehonite. Unlike salt lakes in many other areas of the world, halite, gypsum, and calcite are relatively rare endogenic precipitates in the Great Plains lakes. The detrital fraction of the lacustrine sediments is normally dominated by clay minerals, carbonate minerals, quartz, and feldspars.
Sediment accumulation in these salt lakes is controlled and modified by a wide variety of physical, chemical, and biological processes. Although the details of these modern sedimentary processes can be exceedingly complex and difficult to discuss in isolation, in broad terms, the processes operating in the salt lakes of the Great Plains are ultimately controlled by three basic factors or conditions of the basin: (a) basin morphology; (b) basin hydrology; and (c) water salinity and composition. Combinations of these parameters interact to control nearly all aspects of modern sedimentation in these salt lakes and give rise to four 'end member' types of modern saline lacustrine settings in the Great Plains: (a) clastics-dominated playas; (b) salt-dominated playas; (c) deep water, non-stratified lakes; and (d) deep water, "permanently" stratified lakes.
PMCID: PMC1315329  PMID: 16297237
5.  Evolutionary Origin and Phylogeography of the Diploid Obligate Parthenogen Artemia parthenogenetica (Branchiopoda: Anostraca) 
PLoS ONE  2010;5(8):e11932.
Understanding the evolutionary origin and the phylogeographic patterns of asexual taxa can shed light on the origin and maintenance of sexual reproduction. We assessed the geographic origin, genetic diversity, and phylogeographic history of obligate parthenogen diploid Artemia parthenogenetica populations, a widespread halophilic crustacean.
Methodology/Principal Findings
We analysed a partial sequence of the Cytochrome c Oxidase Subunit I mitochondrial gene from an extensive set of localities (including Eurasia, Africa, and Australia), and examined their phylogeographic patterns and the phylogenetic relationships of diploid A. parthenogenetica and its closest sexual relatives. Populations displayed an extremely low level of mitochondrial genetic diversity, with one widespread haplotype shared by over 79% of individuals analysed. Phylogenetic and phylogeographic analyses indicated a multiple and recent evolutionary origin of diploid A. parthenogenetica, and strongly suggested that the geographic origin of parthenogenesis in Artemia was in Central Asia. Our results indicate that the maternal sexual ancestors of diploid A. parthenogenetica were an undescribed species from Kazakhstan and A. urmiana.
We found evidence for multiple origin of parthenogenesis in Central Asia. Our results indicated that, shortly after its origin, diploid A. parthenogenetica populations underwent a rapid range expansion from Central Asia towards the Mediterranean region, and probably to the rest of its current geographic distribution. This contrasts with the restricted geographic distribution, strong genetic structure, and regional endemism of sexual Artemia lineages and other passively dispersed sexual continental aquatic invertebrates. We hypothesize that diploid parthenogens might have reached their current distribution in historical times, with a range expansion possibly facilitated by an increased availability of suitable habitat provided by anthropogenic activities, such as the spread of solar saltworks, aided by their natural dispersal vectors (i.e., waterbirds).
PMCID: PMC2915914  PMID: 20694140
6.  A biometric and ecologic comparison between Artemia from Mexico and Chile 
Saline Systems  2006;2:13.
A preliminary biometric and ecologic database for the brine shrimp Artemia from Mexico and Chile is presented. The area abounds in small and seasonal ponds and large inland lakes, the latter mainly located in Mexico, although relatively large and isolated lakes are found in complex hydrological settings in pre-high plateau areas of Chile. This paper summarizes research efforts aimed at the localization, characterization, and evaluation of the aquaculture potential of Artemia populations in Mexico and Chile, which exhibit great habitat diversity (ponds, salterns, coastal lagoons, sea arms, coastal and inland lakes), contrasting weather conditions and different levels of isolation and human intervention.
This study covered locations between 29° north latitude (Baja California, Mexico) to 50° south latitude (Puerto Natales, Chile). Biological characteristics considered are species name, reproductive mode, cyst diameter, chorion thickness, and nauplius length, whereas ecological data include pond size, pH, salinity, temperature, and water ionic composition. Artemia franciscana is the only species found in Mexico, it exists together with A. persimilis in Chile, though separated geographically. Ecological differences in habitat exist between both regions but also within countries, a pattern particularly clear with regard to water composition. Surprisingly, a Mexican (Cuatro Ciénegas, A. franciscana) and a Chilean location (Torres del Paine, A. persimilis) share habitat characteristics, at least for the period when data were collected. The discriminat analysis for cyst diameter and nauplius length shows that Artemia from only one location match in cyst diameter with those from San Francisco Bay (SFB) (Point Lobos), and one (Marquez) is far apart from SFB and all the others. The Chilean locations (Pampilla, Cejar, Cahuil, Llamara, Yape) share cyst diameter, but tend to differ from SFB. The remaining Mexican locations (Juchitan, Ohuira, Yavaros) are well separated from all the others. With regard to nauplii length, populations tend to distribute in a relatively random manner, being Marquez the location differing the most in cyst diameter from SFB.
This database will contribute to the knowledge of radiation centers and serves as a baseline for further biogeographic studies, population characterization, management, and monitoring of Artemia biodiversity. Likewise, the impact of colonization and translocations for aquaculture purposes can be better assessed with a baseline for reference. Mexico and Chile exemplify the need to increase and further integrate regional information to tackle fundamental problems underlying practical utilization of Artemia.
PMCID: PMC1687188  PMID: 17125517
7.  Identification and Characterization of Bacteria in a Selenium-Contaminated Hypersaline Evaporation Pond 
Solar evaporation ponds are commonly used to reduce the volume of seleniferous agricultural drainage water in the San Joaquin Valley, Calif. These hypersaline ponds pose an environmental health hazard because they are heavily contaminated with selenium (Se), mainly in the form of selenate. Se in the ponds may be removed by microbial Se volatilization, a bioremediation process whereby toxic, bioavailable selenate is converted to relatively nontoxic dimethylselenide gas. In order to identify microbes that may be used for Se bioremediation, a 16S ribosomal DNA phylogenetic analysis of an aerobic hypersaline pond in the San Joaquin Valley showed that a previously unaffiliated group of uncultured bacteria (belonging to the order Cytophagales) was dominant, followed by a group of cultured γ-Proteobacteria which was closely related to Halomonas species. Se K-edge X-ray absorption spectroscopy of selenate-treated bacterial isolates showed that they accumulated a mixture of predominantly selenate and a selenomethionine-like species, consistent with the idea that selenate was assimilated via the S assimilation pathway. One of these bacterial isolates (Halomonas-like strain MPD-51) was the best candidate for the bioremediation of hypersaline evaporation ponds contaminated with high Se concentrations because it tolerated 2 M selenate and 32.5% NaCl, grew rapidly in media containing selenate, and accumulated and volatilized Se at high rates (1.65 μg of Se g of protein−1 h−1), compared to other cultured bacterial isolates.
PMCID: PMC93092  PMID: 11525968
8.  Molecular identification of microorganisms associated with the brine shrimp Artemia franciscana 
Aquatic Biosystems  2013;9:7.
Prior research on the microorganisms associated with the brine shrimp, Artemia franciscana, has mainly been limited to culture-based identification techniques or feeding studies for aquaculture. Our objective was to identify bacteria and archaea associated with Artemia adults and encysted embryos to understand the role of microbes in the Artemia life cycle and, therefore, their importance in a hypersaline food chain.
We used small subunit (SSU) 16S ribosomal RNA gene sequencing to identify bacteria and archaea associated with adults and encysted Artemia embryos from one of their natural environments – Great Salt Lake (GSL), Utah, USA. We found that bacterial sequences most closely related to the genera Halomonas and Vibrio were commonly extracted from GSL adult Artemia, while bacterial sequences most similar to the genera Halomonas, Psychroflexus and Alkalilimnicola dominate in GSL water. Encysted embryos (cysts) yielded bacterial sequences from the genera Idiomarina and Salinivibrio, which were absent from adults and water. Common archaeal sequences in adults were most closely related to the genera Haloterrigena and Haloarcula, while all of the archaeal sequences from GSL water were most similar to the genus Halogeometricum. Cyst derived archaeal sequences were most closely related to the genera Halorubrum and Haloarcula.
In addition to identifying microbial rRNA sequences that are specific to different stages of the Artemia life cycle, we observed striking differences in the sequences associated with the adult Artemia population in samples collected from GSL at different times and locations. While our study was limited in scope and the sample was small, our findings provide a foundation for future research into how the bacteria and archaea associated with Artemia influence the Artemia life cycle, and GSL food web.
PMCID: PMC3599907  PMID: 23497541
9.  Egg banks in hypersaline lakes of the South-East Europe 
Saline Systems  2009;5:3.
The cyst banks of 6 coastal hypersaline lakes of South-East Europe have been investigated. The study concerned the bottom sediments of Khersonesskoe and Koyashskoe lakes in the Crimea (Ukraine), Nartë saltworks (Albania), Vecchia Salina at Torre Colimena (Apulia, Italy), Pantano Grande and Pantano Roveto at Vendicari (Sicily, Italy). A total of 19 cyst types were recognised. The cyst banks of lakes were found to be well separated in the representation derived from a statistical multivariate data analysis. For all the lakes examined a comparison was possible between the resting community in sediments (cyst bank) and the active one in the water. The cyst banks contained more species than those recorded over a multi-year sampling effort in the water column. The study of cyst hatching, performed on 5 cyst types under lab conditions, demonstrated that cysts do not hatch under the same conditions. Furthermore, each cyst type shows a wide range of preferential hatching conditions, which allow us to confirm the ecological generalism of salt lake species.
PMCID: PMC2662865  PMID: 19292906
10.  A traditional Japanese-style salt field is a niche for haloarchaeal strains that can survive in 0.5% salt solution 
Saline Systems  2007;3:2.
Most of the haloarchaeal strains have been isolated from hypersaline environments such as solar evaporation ponds, salt lakes, or salt deposits, and they, with some exceptions, lyse or lose viability in very low-salt concentrations. There are no salty environments suitable for the growth of haloarchaea in Japan. Although Natrialba asiatica and Haloarcula japonica were isolated many years ago, the question, "Are haloarchaea really thriving in natural environments of Japan?" has remained unanswered.
Ten strains were isolated from a traditional Japanese-style salt field at Nie, Noto Peninsula, Japan by plating out the soil samples directly on agar plates containing 30% (w/v) salts and 0.5% yeast extract. They were most closely related to strains of three genera, Haladaptatus, Halococcus, and Halogeometricum. Survival rates in 3% and 0.5% SW (Salt Water, solutions containing salts in approximately the same proportions as found in seawater) solutions at 37°C differed considerably depending on the strains. Two strains belonging to Halogeometricum as well as the type strain Hgm. borinquense died and lysed immediately after suspension. Five strains that belonged to Halococcus and a strain that may be a member of Halogeometricum survived for 1–2 days in 0.5% SW solution. Two strains most closely related to Haladaptatus possessed extraordinary strong tolerance to low salt conditions. About 20 to 34% of the cells remained viable in 0.5% SW after 9 days incubation.
In this study we have demonstrated that haloarchaea are really thriving in the soil of Japanese-style salt field. The haloarchaeal cells, particularly the fragile strains are suggested to survive in the micropores of smaller size silt fraction, one of the components of soil. The inside of the silt particles is filled with concentrated salt solution and kept intact even upon suspension in rainwater. Possible origins of the haloarchaea isolated in this study are discussed.
PMCID: PMC1828056  PMID: 17346353
11.  Determination of biological and physicochemical parameters of Artemia franciscana strains in hypersaline environments for aquaculture in the Colombian Caribbean 
Saline Systems  2005;1:9.
Artemia (Crustacea, Anostraca), also known as brine shrimp, are typical inhabitants of extreme environments. These hypersaline environments vary considerably in their physicochemical composition, and even their climatic conditions and elevation. Several thalassohaline (marine) environments along the Colombian Caribbean coast were surveyed in order to contribute to the knowledge of brine shrimp biotopes in South America by determining some vital biological and physicochemical parameters for Artemia survival. Additionally, cyst quality tests, biometrical and essential fatty acids analysis were performed to evaluate the economic viability of some of these strains for the aquaculture industry.
In addition to the three locations (Galerazamba, Manaure, and Pozos Colorados) reported in the literature three decades ago in the Colombian Caribbean, six new locations were registered (Salina Cero, Kangaru, Tayrona, Bahía Hondita, Warrego and Pusheo). All habitats sampled showed that chloride was the prevailing anion, as expected, because of their thalassohaline origin. There were significant differences in cyst diameter grouping strains in the following manner according to this parameter: 1) San Francisco Bay (SFB-Control, USA), 2) Galerazamba and Tayrona, 3) Kangarú, 4) Manaure, and 5) Salina Cero and Pozos Colorados. Chorion thickness values were smaller in Tayrona, followed by Salina Cero, Galerazamba, Manaure, SFB, Kangarú and Pozos Colorados. There were significant differences in naupliar size, grouping strains as follows (smallest to largest): 1) Galerazamba, 2) Manaure, 3) SFB, Kangarú, and Salina Cero, 4) Pozos Colorados, and 5) Tayrona. Overall, cyst quality analysis conducted on samples from Manaure, Galerazamba, and Salina Cero revealed that all sites exhibited a relatively high number of cysts.g-1. Essential fatty acids (EFA) analysis performed on nauplii from cyst samples from Manaure, Galerazamba, Salina Cero and Tayrona revealed that cysts from all sites exhibited high arachidonic acid:20:4(n-6) (ArA) and eicosapentaenoic acid: 20:5(n-3) (EPA) levels comparable to the control sample (SFB). In contrast, most cysts collected (including SFB) at different locations, and during different months, presented low docosahexaenoic acid: 22:6(n-3) (DHA) levels (Manaure was the only exception with high DHA levels). Some variations in EPA and ArA levels were observed in all sites, contrasting with the much lower DHA levels which remained constant for all locations, except for Manaure which exhibited variable DHA levels. DHA/EPA ratio was overall very low for all sites compared to SFB cysts. All strains had a low DHA/ArA, but a high EPA/ArA ratio, including the control.
The Colombian A. franciscana habitats analyzed were determined to be thalassohaline, and suitable for A. franciscana development. EFA profiles demonstrated that Tayrona, Galerazamba, Manaure and Salina Cero strains are suitable food for marine fish and crustacean culture because of their high EPA/ArA ratio, but might have to be fortified with DHA rich emulsions depending on the nutritional requirements of the species to be cultured, because of their overall low DHA content. The relatively small nauplii are appropriate for marine larvaeculture. In contrast, the strains from Tayrona, Kangarú, Salina Cero, and Pozos Colorados may be of use but limited to Artemia small biomass production quantities, because of the small surface area of their respective locations; Artemia could be exploited at these locations for local aquaculture applications. In general, cyst quality evaluation for Manaure, Salina Cero and Galerazamba cysts revealed that cysts from these three locations could improve their quality by concentrating efforts on cyst processing techniques. Finally, most locations had great A. franciscana production potential and require different degrees of water quality and/or infrastructure management.
PMCID: PMC1280932  PMID: 16250916
12.  Biodiversity of Archaea and floral of two inland saltern ecosystems in the Alto Vinalopó Valley, Spain 
Saline Systems  2010;6:10.
The extraction of salt from seawater by means of coastal solar salterns is a very well-described process. Moreover, the characterization of these environments from ecological, biochemical and microbiological perspectives has become a key focus for many research groups all over the world over the last 20 years. In countries such as Spain, there are several examples of coastal solar salterns (mainly on the Mediterranean coast) and inland solar salterns, from which sodium chloride is obtained for human consumption. However, studies focused on the characterization of inland solar salterns are scarce and both the archaeal diversity and the plant communities inhabiting these environments remain poorly described.
Two of the inland solar salterns (termed Redonda and Penalva), located in the Alto Vinalopó Valley (Alicante, Spain), were characterized regarding their geological and physico-chemical characteristics and their archaeal and botanical biodiversity. A preliminary eukaryotic diversity survey was also performed using saline water. The chemical characterization of the brine has revealed that the salted groundwater extracted to fill these inland solar salterns is thalassohaline. The plant communities living in this environment are dominated by Sarcocornia fruticosa (L.) A.J. Scott, Arthrocnemum macrostachyum (Moris) K. Koch, Suaeda vera Forsk. ex Gmelin (Amaranthaceae) and several species of Limonium (Mill) and Tamarix (L). Archaeal diversity was analyzed and compared by polymerase chain reaction (PCR)-based molecular phylogenetic techniques. Most of the sequences recovered from environmental DNA samples are affiliated with haloarchaeal genera such as Haloarcula, Halorubrum, Haloquadratum and Halobacterium, and with an unclassified member of the Halobacteriaceae. The eukaryote Dunaliella was also present in the samples.
To our knowledge, this study constitutes the first analysis centered on inland solar salterns located in the southeastern region of Spain. The results obtained revealed that the salt deposits of this region have marine origins. Plant communities typical of salt marshes are present in this ecosystem and members of the Halobacteriaceae family can be easily detected in the microbial populations of these habitats. Possible origins of the haloarchaea detected in this study are discussed.
PMCID: PMC2984398  PMID: 20942947
13.  Community solar salt production in Goa, India 
Aquatic Biosystems  2012;8:30.
Traditional salt farming in Goa, India has been practised for the past 1,500 years by a few communities. Goa’s riverine estuaries, easy access to sea water and favourable climatic conditions makes salt production attractive during summer. Salt produced through this natural evaporation process also played an important role in the economy of Goa even during the Portuguese rule as salt was the chief export commodity. In the past there were 36 villages involved in salt production, which is now reduced to 9. Low income, lack of skilled labour, competition from industrially produced salt, losses incurred on the yearly damage of embankments are the major reasons responsible for the reduction in the number of salt pans.
Salt pans (Mithagar or Mithache agor) form a part of the reclaimed waterlogged khazan lands, which are also utilised for aquaculture, pisciculture and agriculture. Salt pans in Goa experience three phases namely, the ceased phase during monsoon period of June to October, preparatory phase from December to January, and salt harvesting phase, from February to June. After the monsoons, the salt pans are prepared manually for salt production. During high tide, an influx of sea water occurs, which enters the reservoir pans through sluice gates. The sea water after 1–2 days on attaining a salinity of approximately 5ºBé, is released into the evaporator pans and kept till it attains a salinity of 23 - 25ºBé. The brine is then released to crystallizer pans, where the salt crystallises out 25 - 27ºBé and is then harvested.
Salt pans form a unique ecosystem where succession of different organisms with varying environmental conditions occurs. Organisms ranging from bacteria, archaea to fungi, algae, etc., are known to colonise salt pans and may influence the quality of salt produced.
The aim of this review is to describe salt farming in Goa’s history, importance of salt production as a community activity, traditional method of salt production and the biota associated with salt pans.
PMCID: PMC3543363  PMID: 23198813
Salt pan; Goa; Estuary; Community; Khazan; Tidal influx; India; Salt production
14.  Growth Potential of Halophilic Bacteria Isolated from Solar Salt Environments: Carbon Sources and Salt Requirements 
Eighteen strains of extremely halophilic bacteria and three strains of moderately halophilic bacteria were isolated from four different solar salt environments. Growth tests on carbohydrates, low-molecular-weight carboxylic acids, and complex medium demonstrated that the moderate halophiles and strains of the extreme halophiles Haloarcula and Halococcus grew on most of the substrates tested. Among the Halobacterium isolates were several metabolic groups: strains that grew on a broad range of substrates and strains that were essentially confined to either amino acid (peptone) or carbohydrate oxidation. One strain (WS-4) only grew well on pyruvate and acetate. Most strains of extreme halophiles grew by anaerobic fermentation and possibly by nitrate reduction. Tests of growth potential in natural saltern brines demonstrated that none of the halobacteria grew well in brines which harbor the densest populations of these bacteria in solar salterns. All grew best in brines which were unsaturated with NaCl. The high concentrations of Na+ and Mg2+ found in saltern crystallizer brines limited bacterial growth, but the concentrations of K+ found in these brines had little effect. MgSO4 was relatively more inhibitory to the extreme halophiles than was MgCl2, but the reverse was true for the moderate halophiles.
PMCID: PMC241517  PMID: 16346609
15.  Infaunal macrobenthic community dynamics in a manipulated hyperhaline ecosystem: a long-term study 
Aquatic Biosystems  2013;9:20.
Understanding the responses of ecological communities to human-induced perturbations is crucial for establishing conservation goals. Ecological communities are dynamic entities undergoing fluctuations due to their intrinsic characteristics as well as anthropogenic pressures varying over time. In this respect, long-term studies, based on large spatial and temporal datasets, may provide useful information in understanding patterns and processes influencing the communities’ structure. Theoretical evidence suggests that a role of biodiversity is acting as a compensatory buffer against environmental variability by decreasing the temporal variance in ecosystem functioning and by raising the level of community response to perturbations through the selection of better performing species. Therefore, the spatial and temporal changes in the specialization of the community components may be used as an effective tool to monitor the effects of natural and anthropogenic alterations of the environment in dynamic systems. We examined the temporal dynamics of macroinvertebrate community structure in the hyperhaline habitat of Tarquinia Saltworks (central Italy). We aimed at: (i) investigating the relationships between the level of community specialization and the alterations of the environment across fourteen years; (ii) comparing the ability of aggregate community parameters such as the average abundance vs. species specialization in describing patterns of community composition.
We arranged the data in three sub-sets according to three periods, each characterized by different environmental conditions. The mean abundance of sampled macroinvertebrates showed a significant change (p < 0.01) only in the community inhabiting the saltwork basin closely connected to the sea, characterized by the highest environmental variation (i.e. the coefficient of variation, CV, of the aggregate environmental variability over the study period, CVrange = 0.010 - 0.2). Here we found marine species like Modiolus adriaticus (Lamarck, 1819), Neanthes irrorata (Malmgren, 1867), and Amphiglena mediterranea (Leydig, 1851), which inhabited the saltworks during the halt period but disappeared during the subsequent eutrophication phase. Conversely, species specialization showed a significant decrease for each sampled community in the presence of habitat degradation and a recovery after ecological restoration. The widest fluctuations of specialization were recorded for the community inhabiting the saltwork basin with the highest long-term environmental variability.
Recent advances have shown how the increased temporal and spatial variability of species’ abundance within the communities may be a signal of habitat disturbance, even in the absence of an apparent decline. Such approach could also be used as a sensitive monitoring tool, able to detect whether or not communities are subjected to increasing biotic homogenization. Also, the increased functional similarity triggered by habitat degradation may impact on species at higher trophic levels, such as the waterbirds wintering in the area or using it as a stopover during migration.
PMCID: PMC4175097  PMID: 24192133
Hyperhaline habitat; Wetlands; Central Tyrrhenian Sea; Biomonitoring; Macroinvertebrates community; Long-term study; Species specialization index; Biodiversity homogenization
16.  Inland hypersaline lakes and the brine shrimp Artemia as simple models for biodiversity analysis at the population level 
Saline Systems  2006;2:14.
Biodiversity can be measured at different hierarchical levels, from genetic diversity within species to diversity of ecosystems, though policy-makers tend to use species richness. The 2010 goal of reducing biodiversity loss, agreed by the subscribers to the Convention on Biological Diversity, requires simple and reliable protocols to evaluate biodiversity at any level in a given ecosystem. Stakeholders, particularly policy makers, need to understand how ecosystem components interact to produce social and economic benefits on the long run, whilst scientists are expected to fulfil this demand by testing and modelling ideally simple (low diversity) ecosystems, and by monitoring key species. This work emphasizes the unique opportunity offered by inland, isolated salt lakes and the brine shrimp Artemia, an example of biodiversity contained at the intra-specific level, as simple models to understand and monitor biodiversity, as well as to assess its predicted positive association with ecosystem stability. In addition to having well identified species and strains and even clones, that allow to test reproductive effects (sexual versus asexual), Artemia benefits from the possibility to set up experimental testing at both laboratory scale and outdoor pond systems, for which a comprehensive cyst bank with sufficient amount of samples from all over the world is available.
PMCID: PMC1684253  PMID: 17132175
17.  Microbial Diversity in Maras Salterns, a Hypersaline Environment in the Peruvian Andes 
Maras salterns are located 3,380 m above sea level in the Peruvian Andes. These salterns consist of more than 3,000 little ponds which are not interconnected and act as crystallizers where salt precipitates. These ponds are fed by hypersaline spring water rich in sodium and chloride. The microbiota inhabiting these salterns was examined by fluorescence in situ hybridization (FISH), 16S rRNA gene clone library analysis, and cultivation techniques. The total counts per milliliter in the ponds were around 2 × 106 to 3 × 106 cells/ml, while the spring water contained less than 100 cells/ml and did not yield any detectable FISH signal. The microbiota inhabiting the ponds was dominated (80 to 86% of the total counts) by Archaea, while Bacteria accounted for 10 to 13% of the 4′,6′-diamidino-2-phenylindole (DAPI) counts. A total of 239 16S rRNA gene clones were analyzed (132 Archaea clones and 107 Bacteria clones). According to the clone libraries, the archaeal assemblage was dominated by microorganisms related to the cosmopolitan square archaeon “Haloquadra walsbyi,” although a substantial number of the sequences in the libraries (31% of the 16S rRNA gene archaeal clones) were related to Halobacterium sp., which is not normally found in clone libraries from solar salterns. All the bacterial clones were closely related to each other and to the γ-proteobacterium “Pseudomonas halophila” DSM 3050. FISH analysis with a probe specific for this bacterial assemblage revealed that it accounted for 69 to 76% of the total bacterial counts detected with a Bacteria-specific probe. When pond water was used to inoculate solid media containing 25% total salts, both extremely halophilic Archaea and Bacteria were isolated. Archaeal isolates were not related to the isolates in clone libraries, although several bacterial isolates were very closely related to the “P. halophila” cluster found in the libraries. As observed for other hypersaline environments, extremely halophilic bacteria that had ecological relevance seemed to be easier to culture than their archaeal counterparts.
PMCID: PMC1489619  PMID: 16751493
18.  Effect of salinity stress on the life history variables of Branchipus schaefferi Fisher, 1834 (Crustacea: Anostraca) 
Saline Systems  2005;1:4.
Freshwater anostracans inhabit ephemeral water bodies in which as the water level decreases due to evaporation the salt concentration increases. Thus, for most anostracans salinity becomes the major stress factor.
We tested five concentrations of NaCl (0 to 8 g/l) on the life table demography of Branchipus schaefferi fed Chlorella (alga). Age-specific survivorship curves of male and female B. schaefferi showed nearly a similar pattern in that increased salt concentration resulted in decreased survivorship. The age-specific reproduction (mx) of females showed several peaks of cyst production at 0 and 1 g/l salinity while in treatments containing salt at 4 or 8 g/l, there were fewer peaks. Average lifespan, life expectancy at birth, gross and net reproductive rates, generation time and the rate of population increase were all significantly influenced by the salt concentration in the medium. The highest value of net reproductive rate (970 cysts/female) was in treatments containing 0 g/l of salt, while the lowest was 13 cysts/female at 8 g/l. The rate of population increase (r) varied from 0.52 to 0.32 per day depending on the salt concentration in the medium.
The low survival and offspring production of B. schaefferi at higher salinity levels suggests that this species is unlikely to colonize inland saline water bodies. Therefore, the temporary ponds in which it is found, proper conservative measures must be taken to protect this species.
PMCID: PMC1183265  PMID: 16176591
19.  The Genus Prochromadora with a Redescription of P. orleji from a Marine Saltern in the People's Republic of China 
Journal of Nematology  1991;23(4):491-501.
The genus Prochromadora Filipjev, 1922 comprising nine species is reviewed and a key is presented. Salient diagnostic characters of the genus are a single, large, solid dorsal tooth and a cuticle showing transverse rows of homogenous punctations that extend around the body without intervening lateral differentiation. A redescription and drawings of P. orleji from a solar saltworks in Qingdao, Shandong, the People's Republic of China, are presented.
PMCID: PMC2619195  PMID: 19283160
marine nematode; marine saltern; nematode; morphology; People's Republic of China; Prochromadora; P. argentinensis; P. asupplementa; P. bulbosa; P. exigua; P. magna; P. megodonta; P. orleji; P. spitzbergensis; P. trisupplementa
20.  Microbial life at high salt concentrations: phylogenetic and metabolic diversity 
Saline Systems  2008;4:2.
Halophiles are found in all three domains of life. Within the Bacteria we know halophiles within the phyla Cyanobacteria, Proteobacteria, Firmicutes, Actinobacteria, Spirochaetes, and Bacteroidetes. Within the Archaea the most salt-requiring microorganisms are found in the class Halobacteria. Halobacterium and most of its relatives require over 100–150 g/l salt for growth and structural stability. Also within the order Methanococci we encounter halophilic species. Halophiles and non-halophilic relatives are often found together in the phylogenetic tree, and many genera, families and orders have representatives with greatly different salt requirement and tolerance. A few phylogenetically coherent groups consist of halophiles only: the order Halobacteriales, family Halobacteriaceae (Euryarchaeota) and the anaerobic fermentative bacteria of the order Halanaerobiales (Firmicutes). The family Halomonadaceae (Gammaproteobacteria) almost exclusively contains halophiles. Halophilic microorganisms use two strategies to balance their cytoplasm osmotically with their medium. The first involves accumulation of molar concentrations of KCl. This strategy requires adaptation of the intracellular enzymatic machinery, as proteins should maintain their proper conformation and activity at near-saturating salt concentrations. The proteome of such organisms is highly acidic, and most proteins denature when suspended in low salt. Such microorganisms generally cannot survive in low salt media. The second strategy is to exclude salt from the cytoplasm and to synthesize and/or accumulate organic 'compatible' solutes that do not interfere with enzymatic activity. Few adaptations of the cells' proteome are needed, and organisms using the 'organic-solutes-in strategy' often adapt to a surprisingly broad salt concentration range. Most halophilic Bacteria, but also the halophilic methanogenic Archaea use such organic solutes. A variety of such solutes are known, including glycine betaine, ectoine and other amino acid derivatives, sugars and sugar alcohols. The 'high-salt-in strategy' is not limited to the Halobacteriaceae. The Halanaerobiales (Firmicutes) also accumulate salt rather than organic solutes. A third, phylogenetically unrelated organism accumulates KCl: the red extremely halophilic Salinibacter (Bacteroidetes), recently isolated from saltern crystallizer brines. Analysis of its genome showed many points of resemblance with the Halobacteriaceae, probably resulting from extensive horizontal gene transfer. The case of Salinibacter shows that more unusual types of halophiles may be waiting to be discovered.
PMCID: PMC2329653  PMID: 18412960
21.  Patterns of microbial diversity along a salinity gradient in the Guerrero Negro solar saltern, Baja CA Sur, Mexico 
The goal of this study was to use environmental sequencing of 16S rRNA and bop genes to compare the diversity of planktonic bacteria and archaea across ponds with increasing salinity in the Exportadora de Sal (ESSA) evaporative saltern in Guerrero Negro, Baja CA S., Mexico. We hypothesized that diverse communities of heterotrophic bacteria and archaea would be found in the ESSA ponds, but that bacterial diversity would decrease relative to archaea at the highest salinities. Archaeal 16S rRNA diversity was higher in Ponds 11 and 12 (370 and 380 g l−1 total salts, respectively) compared to Pond 9 (180 g l−1 total salts). Both Pond 11 and 12 communities had high representation (47 and 45% of clones, respectively) by Haloquadratum walsbyi-like (99% similarity) lineages. The archaeal community in Pond 9 was dominated (79%) by a single uncultured phylotype with 99% similarity to sequences recovered from the Sfax saltern in Tunisia. This pattern was mirrored in bop gene diversity with greater numbers of highly supported phylotypes including many Haloquadratum-like sequences from the two highest salinity ponds. In Pond 9, most bop sequences, were not closely related to sequences in databases. Bacterial 16S rRNA diversity was higher than archaeal in both Pond 9 and Pond 12 samples, but not Pond 11, where a non-Salinibacter lineage within the Bacteroidetes >98% similar to environmental clones recovered from Lake Tuz in Turkey and a saltern in Chula Vista, CA was most abundant (69% of community). This OTU was also the most abundant in Pond 12, but only represented 14% of clones in the more diverse pond. The most abundant OTU in Pond 9 (33% of community) was 99% similar to an uncultured gammaproteobacterial clone from the Salton Sea. Results suggest that the communities of saltern bacteria and archaea vary even in ponds with similar salinity and further investigation into the ecology of diverse, uncultured halophile communities is warranted.
PMCID: PMC3868825  PMID: 24391633
halophile; gradient; saltern; 16S rRNA gene; bop gene; haloarchaea
22.  Artisanal salt production in Aveiro/Portugal - an ecofriendly process 
Saline Systems  2011;7:3.
Solar salinas are man-made systems exploited for the extraction of salt, by solar and wind evaporation of seawater. Salt production achieved by traditional methods is associated with landscapes and environmental and patrimonial values generated throughout history. Since the mid-twentieth century, this activity has been facing a marked decline in Portugal, with most salinas either abandoned or subjected to destruction, making it necessary to find a strategy to reverse this trend.
It is, however, possible to generate revenue from salinas at several levels, not merely in terms of good quality salt production, but also by obtaining other products that can be commercialized, or by exploring their potential for tourism, and as research facilities, among others. Furthermore, with an adequate management, biodiversity can be restored to abandoned salinas, which constitute important feeding and breeding grounds for resident and migratory aquatic birds, many of which are protected by European Community Directives.
The aims of this manuscript are to present a brief overview on the current state of sea salt exploitation in Portugal and to stress the importance of recovering these salinas for the conservation of this particular environment, for the regional economy, the scientific community and the general public. The Aveiro salina complex is presented in detail, to exemplify salina structure and functioning, as well as current problems and potential solutions for artisanal salinas.
PMCID: PMC3225316  PMID: 22053788
Salt production; Salinas; Coastal wetlands; Biodiversity; Ecosystems; Aveiro; Portugal
23.  Phylogenetic diversity of actinomycetes cultured from coastal multipond solar saltern in Tuticorin, India 
Aquatic Biosystems  2012;8:23.
Hypersaline solar salterns are extreme environments in many tropical and subtropical regions throughout the world. In India, there are several coastal solar salterns along with the coastal line of the Bay of Bengal and Arabian Sea and inland solar salterns around Sambhar saltlake, from which sodium chloride is obtained for human consumption and industrial needs. Studies on characterization of such coastal and inland solar salterns are scarce and both the bacterial and archaeal diversity of these extreme saline environment remains poorly understood. Moreover, there are no reports on exclusive diversity of actinomycetes inhabiting Indian solar salterns.
Soil sediments were collected from both concentrator and crystallizer ponds of solar salterns and subjected to detailed physico-chemical analysis. Actinomycetes were selectively isolated by employing selective processing methods and agar media. A total of 12 representatives were selected from the 69 actinomycete isolates obtained from the saltern soil samples, using Amplified Ribosomal DNA Restriction Analysis. Sequencing and analysis of 16S rDNA from chosen representative isolates displayed the presence of members affiliated to actinobacterial genera: Streptomyces, Micromonospora, Nocardia, Nocardiopsis, Saccharopolyspora and Nonomuraea. The genus Streptomyces was found to be the dominant among the isolates. Furthermore, rare actinomycete genus Nonomuraea was isolated for the first time from Indian solar salterns.
To the best of our knowledge, this study constitutes the first characterization of actinomycete diversity centred on solar salterns located in the eastern coastal region of India. Furthermore, this is the very first report of isolation of Nonomuraea species from solar salterns and also from India. As actinomycetes encompass recurrently foremost sources of biotechnologically important member of the microbial communities, the actinomycetes retrieved from the Indian saltern soil samples laid the platform to search for novel biotechnologically significant bioactive substances.
PMCID: PMC3496644  PMID: 22950748
Solar saltern; Actinomycete; ARDRA; Phylogeny; 16S rDNA; Nonomuraea
24.  Ecology and ethnoecology of dusky grouper [garoupa, Epinephelus marginatus (Lowe, 1834)] along the coast of Brazil 
Dusky grouper (garoupa, Epinephelus marginatus) is an important catch for several artisanal small-scale fisheries along the Brazilian coast. It is a sedentary, monandric, and late maturing protogynous species, which makes it vulnerable to overharvesting even though it is mainly caught through hook and line or spear fishing through free diving. Lack of information on the ecology and biology of this species in Brazil is astonishing. Much of the information found in the literature concerns Mediterranean dusky groupers. Studies compiling local knowledge (ethnoecology) about fish species complement biological data, and have been fundamental for effective fisheries management. In this study, our objectives are to obtain data about dusky grouper through fish catches and analysis of stomach contents and gonad maturation (macroscopic analyses), along with interviews from fishermen from six small-scales communities from the southern (Pântano do Sul, Florianópolis, Santa Catarina State) to the northern Brazilian coast (Porto Sauípe, Bahia State). We conclude that precautionary approaches and 'data-less' management approaches are needed in the coast of Brazil. Research on this species and on the potential of aquaculture for its cultivation, are urgent, due to the apparent vulnerability and decrease of dusky grouper along the coast of Brazil.
PMCID: PMC2567293  PMID: 18793394
25.  Optimization studies on production of a salt-tolerant protease from Pseudomonas aeruginosa strain BC1 and its application on tannery saline wastewater treatment 
Brazilian Journal of Microbiology  2011;42(4):1506-1515.
Treatment and safe disposal of tannery saline wastewater, a primary effluent stream that is generated by soaking salt-laden hides and skin is one of the major problems faced by the leather manufacturing industries. Conventional treatment methods like solar evaporation ponds and land composting are not eco-friendly as they deteriorate the ground water quality. Though, this waste stream is comprised of high concentration of dissolved proteins the presence of high salinity (1–6 % NaCl by wt) makes it non-biodegradable. Enzymatic treatment is one of the positive alternatives for management of such kind of waste streams. A novel salt-tolerant alkaline protease obtained from P.aeruginosa (isolated from tannery saline wastewater) was used for enzymatic degradation studies. The effect of various physical factors including pH, temperature, incubation time, protein source and salinity on the activity of identified protease were investigated. Kinetic parameters (Km , Vmax) were calculated for the identified alkaline protease at varying substrate concentrations. Tannery saline wastewater treated with identified salt tolerant protease showed 75 % protein removal at 6 h duration and 2 % (v/v) protease addition was found to be the optimum dosage value.
PMCID: PMC3768711  PMID: 24031785
COD; Enzymatic degradation; Leather Industry; Salt-tolerant; Waste treatment

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