Relationships between Artemia monica life history characteristics and salinity

Relationships between Artemia monica life history characteristics and salinity were determined using data from four published studies and three experiments presented here. Salinity explained 40 to 93 percent of the variation in ten life history characteristics. Reductions in hatching success, survival, length, weight, ovigery, and brood size were observed as salinity increased from 76 to 168 g l^–1. Inter-brood duration, and time to hatching and reproduction were protracted as salinity was elevated. Salinity effects on life history characteristics appeared to be gradual and continuous rather than exhibiting thresholds. The one exception was naupliar survival, which was constant between 76 and 133 g l^–1 followed by a decrease above 133 g l^–1.     

Anostracan cysts found in California salt lakes

The salt lakes of California are home to four genera and eight species of Anostraca. The cysts of most of these species are readily distinguished using a combination of cyst and geographic characters. Cyst identification is a valuable tool that can allow a look at fairy shrimp ecology when adults are not available. For example, fossil anostracan cysts from 500-year-old Mono Lake sediments show more character variation than currently exists in extant Artemia monica, indicating a greater diversity of Anostraca at that time. Continuing desertification, both natural and man-made, in the Mono Lake area over the last 500 years is associated with decreased diversity in the Mono Lake anostracan fauna.     

In situ hatching of Artemia monica cysts in hypersaline Mono Lake,California

Two emergence trap designs were tested in Mono Lake, California, to measure in situ hatching of Artemia monica cysts on the lake bottom. One design incorporated a removable sample bottle; the other had a catch tube which was pumped from the surface. Both traps rested on the bottom and had a narrow gap between the collecting funnel and bottom flange to allow the chemical conditions within the trap to be similar to those outside. This gap was open during April and May but, because some animals entered from outside the area enclosed by the trap, the gap was covered with 400 µm or 800 µm screen during June and July. The two trap types without screens sampled a station in oxic water 7 m deep similarly in April and May 1985. Mean daily hatching rates from April to May 1985 ranged from 720 to 25 340 shrimp m^-2 day^-1. In contrast, mean daily hatching rates during the same period at a station in anoxic water 21 m deep were from 3 to 138 shrimp m^-2 day^-1. June and July hatching rates in the shallow station were lower than in the spring, usually less than 1000 shrimp m^-2 day^-1.     

SALINITY AND NUTRIENT LIMITATIONS ON GROWTH OF BENTHIC ALGAE FROM TWO ALKALINE SALT LAKES OF THE WESTERN GREAT BASIN (USA)1

Enrichment cultures of littoral benthic algae from Mono Lake, California, and Abert Lake, Oregon, were grown under conditions of varied salinity and nutrient content. Field-collected inocula were composed mainly of diatoms and filamentous blue-green and green algae. The yield of long-term cultures (30 days) showed tolerance over a broad salinity range (50–150 g·L^?1) for Mono Lake-derived algae. Algae from Abert Lake had a lower range of tolerance (25–100 g·L^?1) Organic content and pigment concentrations of algae from both lakes were also reduced above the tolerated salinity level. Within the range of salinity tolerance for Mono Lake algae, initial growth rates and organic content were reduced by increased salinity. The effects of macro- and micronutrient enrichment on algal growth in Mono Lake water were also tested. Only nitrogen enrichment (either as ammonium or nitrate) stimulated algal growth. Although the benthic algae cultured here had wide optima for salinity tolerance, the rates of growth and storage were limited by increased salinity within the optimum range. Although the lakes compared had similar species composition, the range and limits of tolerance of the algae were related to salinity of the lake of origin.     

Genetic characterization and intra-generic relationships of Artemia monica Verrill and A. urmiana Günther

Allozyme variation encoded for by 22 enzyme loci analyzed with starch-gel electrophoresis from samples of gonochoristic Artemia monica and A. urmiana collected from Lake Urmia, Iran and from Mono Lake, CA, USA, respectively, are compared and contrasted with data from representative gonochoristic populations of A. franciscana, A. persimilis and A. salina, as well as diploid, triploid, tetraploid, and pentaploid parthenogenetic forms. Values for parameters measuring degree of genetic variability (number of alleles per locus, proportion of polymorphic loci, and expected heterozygosity) in A. monica and A. urmiana were among the highest in the genus (1.76, 0.46, 0.19, and 2.19, 0.55, and 0.14, respectively). For A. monica, possible factors promoting its high genetic variability are discussed. Mean values for Nei's D between A. monica and A. franciscana were very low (0.09) and consistent with reported cytological and morphological similarities, thus the physiological differentiation which distinguishes the former must involve differences at very few loci. D values from A. urmiana to rest of the gonochoristic species (0.95) are consistent with its taxonomic status and distinctive morphology, while its low genetic distance to the monophyletic Artemia parthenogenetic lineage (mean D = 0.48) suggests a recent common ancestry.     

Nutrient fluxes from upwelling and enhanced turbulence at the top of the pycnocline in Mono Lake, California

Time series measurements of temperature at 15 depths and profiles of temperature-gradient microstructure were obtained during a period with strong wind forcing and subsequent calm in Mono Lake, California. The wind forcing increased the amplitude of basin-scale internal waves and energy at all wave frequencies relative to the calm period. Rates of dissipation of turbulent kinetic energy, , were high ( > 10^–6 m² s^–3) at the top of the pycnocline at both an inshore and an offshore site on a day when winds reached 10 m s^–1 and on the following two days at an inshore site ( > 10^–7 m² s^–3). The enhanced turbulence occurred at the depth of a subsurface temperature maximum (z _TM) and coincidentally with elevated concentrations of NH₄, reduced concentrations of chlorophyll a and particulate carbon, and increased abundance of the macrozooplankter Artemia monica. The NH₄ at z _TM was more dispersed and of lower concentration inshore than offshore and indicated greater turbulent transport inshore. Over the course of 4 days, chlorophyll a concentrations increased in the upper mixed layer, and C:N and C:Chl ratios decreased. Offshore, the change in C:N ratio indicated a relaxation of moderate nutrient deficiency. We hypothesize that excretion by A. monica and turbulent transport of the NH₄ from the subsurface temperature maximum led to improved physiological status of phytoplankton in the upper mixed layer.     

Effects of Imposed Salinity Gradients on Dissimilatory Arsenate Reduction, Sulfate Reduction, and Other Microbial Processes in Sediments from Two California Soda Lakes

Salinity effects on microbial community structure and on potential rates of arsenate reduction, arsenite oxidation, sulfate reduction, denitrification, and methanogenesis were examined in sediment slurries from two California soda lakes. We conducted experiments with Mono Lake and Searles Lake sediments over a wide range of salt concentrations (25 to 346 g liter⁻¹). With the exception of sulfate reduction, rates of all processes demonstrated an inverse relationship to total salinity. However, each of these processes persisted at low but detectable rates at salt saturation. Denaturing gradient gel electrophoresis analysis of partial 16S rRNA genes amplified from As(V) reduction slurries revealed that distinct microbial populations grew at low (25 to 50 g liter⁻¹), intermediate (100 to 200 g liter⁻¹), and high (>300 g liter⁻¹) salinity. At intermediate and high salinities, a close relative of a cultivated As-respiring halophile was present. These results suggest that organisms adapted to more dilute conditions can remain viable at high salinity and rapidly repopulate the lake during periods of rising lake level. In contrast to As reduction, sulfate reduction in Mono Lake slurries was undetectable at salt saturation. Furthermore, sulfate reduction was excluded from Searles Lake sediments at any salinity despite the presence of abundant sulfate. Sulfate reduction occurred in Searles Lake sediment slurries only following inoculation with Mono Lake sediment, indicating the absence of sulfate-reducing flora. Experiments with borate-amended Mono Lake slurries suggest that the notably high (0.46 molal) concentration of borate in the Searles Lake brine was responsible for the exclusion of sulfate reducers from that ecosystem.     

EXPERIMENTAL MESOCOSM STUDIES OF SALINITY EFFECTS ON THE BENTHIC ALGAL COMMUNITY OF A SALINE LAKE

As closed-basin systems, saline lakes are prone to fluctuate in level and salinity with climate change and hydrologic alterations. Loss of many Great Basin lakes has resulted from the diversion of tributary streams for agricultural or municipal uses. At Mono Lake, an alkaline salt lake in eastern California, salinities have risen from 50 to 100 g·L^?1 in just 50 years. Experimental mesocosms were established to simulate some of the potential ecological effects that could have accompanied this change. The influence of salinity on diatom diversity, taxonomic structure, and primary production was tested using mesocosms deployed at Mono Lake. Mesocosm tanks were 500 L in volume, 1 m square, and 0.5 m deep, with open tops covered by 1 mm mesh net. Five treatments (50, 75, 100, 125, and 150 g·L^?1) with four replicates per treatment were used over a 2-month period. The diatom-dominated benthic algae were reduced both in standing crop (from 6 to <0.1 g·m^?2) and diversity (from 30 to 12 taxa) with increased salinity, with most loss occurring in salinities ≥75 g·L^?1. Photosynthetic oxygen production also was significantly lower at salinities ≥75 g·L^?1. Diatom indicator taxa for these shifts included Denticula sp., Nitzschia frustulum, N. monoensis, N. communis, and Stephanodiscus oregonicus increasing in relative abundance in higher salinity treatments, accompanied by decreases in Achnanthes minutissima, Cymbella minuta, N. dissipata, and Rhoicosphenia abbreviata. Exhibiting dominance at moderate salinity levels (75 to 125 g·L^?1) were Nitzschia frustulum, N. communis, N. palea, and Navicula crucialis. These latter species may be limited by both physiological stress at high salinity and grazing and competition at low salinity. The filamentous chlorophyte, Ctenocladus circinnatus, and cyanobacteria (Oscillatoria spp.) occurred only in salinity treatments from 50 to 100 g·L^?1. Diversion of tributary stream flow and resulting salinity increases in this lake threaten sustained benthic primary production and algal species diversity relative to conditions prior to stream diversion. The 1994 decision of the California State Water Resources Control Board to return stream flows to Mono Lake will raise the lake level and reduce salinity to around 75 g·L^?1 and is expected to increase the diversity and productivity of the benthic algae of this ecosystem.     

Comparative population ecology of Ephydra hians Say (Diptera: Ephydridae) at Mono Lake (California) and Abert Lake (Oregon)

The population dynamics of Ephydra hians Say final instar larvae and pupae were compared over a two year period in rocky littoral habitats of two alkaline saline lakes in the western Great Basin. Relative abundance increased from 1983 to 1984 at Mono Lake (California), during dilution from ca. 90 to 80 g 1^-1 TDS (total dissolved solids). In contrast, relative abundance decreased over the same period at Abert Lake (Oregon), accompanied by a dilution of salinity from ca. 30 to 20 g l^-1 and a marked increase in the number and abundance of other benthic macroinvertebrate species. These observations are consistent with a hypothesis that proposes biotic interactions limit E. hians abundance at low salinity, and physiological stress limits abundance at high salinity.Oviposition extends from early spring to early fall. Mixed instars present throughout this period indicates multivoltine population dynamics with overlapping generations. The standing stock biomass of final instars increases exponentially in late spring and peaks in late summer or early fall. Pupae increase in proportional representation and abundance from a spring minimum to a fall maximum. The body size of adults and pupae cycle seasonally from a spring maximum to a fall minimum, and may be related to either or both food limitation, or water temperature.     

Effects of imposed salinity gradients on dissimilatory arsenate reduction, sulfate reduction, and other microbial processes in sediments from two California soda lakes

Salinity effects on microbial community structure and on potential rates of arsenate reduction, arsenite oxidation, sulfate reduction, denitrification, and methanogenesis were examined in sediment slurries from two California soda lakes. We conducted experiments with Mono Lake and Searles Lake sediments over a wide range of salt concentrations (25 to 346 g liter(-1)). With the exception of sulfate reduction, rates of all processes demonstrated an inverse relationship to total salinity. However, each of these processes persisted at low but detectable rates at salt saturation. Denaturing gradient gel electrophoresis analysis of partial 16S rRNA genes amplified from As(V) reduction slurries revealed that distinct microbial populations grew at low (25 to 50 g liter(-1)), intermediate (100 to 200 g liter(-1)), and high (>300 g liter(-1)) salinity. At intermediate and high salinities, a close relative of a cultivated As-respiring halophile was present. These results suggest that organisms adapted to more dilute conditions can remain viable at high salinity and rapidly repopulate the lake during periods of rising lake level. In contrast to As reduction, sulfate reduction in Mono Lake slurries was undetectable at salt saturation. Furthermore, sulfate reduction was excluded from Searles Lake sediments at any salinity despite the presence of abundant sulfate. Sulfate reduction occurred in Searles Lake sediment slurries only following inoculation with Mono Lake sediment, indicating the absence of sulfate-reducing flora. Experiments with borate-amended Mono Lake slurries suggest that the notably high (0.46 molal) concentration of borate in the Searles Lake brine was responsible for the exclusion of sulfate reducers from that ecosystem.     

GROWTH OF THE FILAMENTOUS GREEN ALGA CTENOCLADUS CIRCINNATUS (CHAETOPHORALES,CHLOROPHYCEAE) IN RELATION TO ENVIRONMENTAL SALINITY1

Clones of the filamentous green alga Ctenocladus circinnatus Borzi were isolated from algae collected at Abert Lake (Oregon) and Mono Lake (California). Stock cultures were exposed to varied salinities of natural lake water to examine the effects on growth rate, cell form, chlorophyll a, and water content. Growth rates were reduced in both clones with increased salinity over the range 25–100 g·L^?1 and were almost completely inhibited at 150 g·L^?1. Chlorophyll a increased between salinities of 25 and 100 g·L^?1, reflecting slower growth, higher proportions of akinetes, and smaller cell sizes as salinity increased. Tissue water content remained essentially constant from 25 to 100 g·L^?1 salinity. Shorter cell dimensions with increased salinity suggest that a lower surface-to-volume ratio may reduce the potential for passive loss of cell water. Prior acclimation of stock cultures to elevated salinity provided no enhancement of growth response at any salinity. The results indicate that environmental salinity can limit the productivity and distribution of Ctenocladus in nature.     

Structure and formation properties of the haloalkaliphilic community of Lake Khilganta

The structural features of a cyanobacterial mat from Lake Khilganta (Southeastern Transbaikal Region) developing at different values of salinity and pH were determined based on our long-term investigation of the natural community, as well as results obtained during experimentation with its laboratory analogue. At water mineralization of 40–50 g/l, Microcoleus chthonoplastes and Phormidium molle play a key role in the formation of the cyanobacterial mat. As water mineralization increases, the diversity of cyanobacteria in the natural mat increases as well, reaches its maximum at 80 g/l NaCl, and decreases at 100 g/l. In the laboratory community, Nodularia sp. prevailed. It was able to form matlike structures within a broad pH range and at a salinity of up to 50 g/l NaCl. As the water mineralization level increased up to 100 g/l or higher, a replacement of the dominant complexes occurred both in the laboratory and natural communities: cyanobacterial species were substituted with green algae.     

Life-history analysis of an Artemia population in a changing environment

The Anemia monica Verrill population in Mono Lake, Californiahas two generations per year. Despite similarities in the year-to-yearlife history patterns, some important differences developedbetween 1979 and 1981. The first generation hatches from overwinteringcysts in early spring and reaches maturity by the end of May.The first-generation females reproduce ovoviviparously, givingrise to a second generation which matures between mid-July andAugust. In July, both first and second generation females beginproducing overwintering cysts. The population reaches it maximumin late summer, then declines to low numbers by November. Theabundance of the first generation in June declined from a meanof 20 000 m^–2 to 2400 m^–2. Despite the smaller firstgeneration, the second generation in 1980 and 1981 was at leastas abundant as in 1979. These differences are indicative ofa change in the Artemia population dynamics in Mono Lake. ¹Address for correspondence: Hawaii Institute of Marine Biology,University of Hawaii, P.O. Box 1346 Kaneohe, HI 96744-1346,USA.     

Nearshore and pelagic abundances of Artemia monica in Mono Lake,California

The spatial distribution and abundance of the brine shrimp, Artemia monica, in Mono Lake, California were determined during 1982 and 1983. Peak abundances of shrimp occur in midsummer and reach densities of 15–17 individuals l^-1 in the nearshore regions and 6–8 individuals 1^-1 in the pelagic region. The brine shrimp were non-uniformly distributed both vertically and horizontally. The coefficient of variation in shrimp abundance among stations within the nearshore region was similar to that found in the pelagic region. On two of the nine dates, nearshore densities were 3 to 4 times greater than those in the pelagic zone, and on average the brine shrimp appear to be slightly over-dispersed to the nearshore region. However, including nearshore abundances in lakewide estimates will usually result in a change of less than a 10%.     

Effect of salinity stress on the life history variables of Branchipus schaefferi Fisher, 1834 (Crustacea: Anostraca)

Background

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.

Results

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 (m_x) 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.

Conclusion

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.     

Spatial and temporal distribution patterns of three trophic levels in a saline lake

Horizontal distribution patterns of three trophic levels wereinvestigated at Mono Lake, California, USA during the autumnsof 1980 and 1981. Coefficients of variation for Anemia monicaand grebes peaked in October of both years, whereas coefficientsof variation for phytoplankton declined from August to November.Stable population gradients were observed in the Anemia population.In contrast, grebes were very mobile and their distributionalpatterns changed between sampling dates. Anemia and phytoplanktonwere negatively correlated during 1980, but no significant correlationsoccurred in 1981. Grebes and Artemia were positively correlatedin October 1980 and 1981, suggesting active prey tracking bythe grebes. Interactions between trophic levels may have contributedto the creation and maintenance of patchiness, although theseeffects appeared to be intermittent.     

Influence of Salinity Concentration on Aquatic Insect Community Structure: A Mesocosm Experiment in the Dead Sea Basin Region

Salinity varies considerably among temporary pools in the Dead Sea Basin, Israel. We experimentally assessed the effects of four salinity levels (0, 10, 20 and 30 g NaCl per liter) on the aquatic insect community in this basin in an artificial pool experiment. Each salinity level was randomly assigned to six pools (total=24 pools). Salinity did not affect total insect abundance but strongly affected abundance and distributions of different species, and consequently, community structure. Of 13 taxa colonizing the pools, 12 were Diptera including 10 mosquito species. Five taxa were sufficiently common to assess abundance in relation to salinity. Polypedilum nubiferum Skuse (Diptera: Chironomidae) was largely salinity intolerant being abundant only in the freshwater. Ephydra flavipes Macquart (Diptera: Ephydridae) was most abundant at the highest salinity level and was rare in freshwater. Ochlerotatus caspius Pallas (Diptera: Culicidae) abundance tended to be highest at 10 g/l and lowest at 30 g/l although the differences were not statistically significant. Anopheles multicolor Cambouliu (Diptera: Culicidae) was relatively euryhaline although numbers dropped significantly at the highest salinity. Cleon dipterum Linnaeus (Baetidae: Ephemeroptera) was also euryhaline and showed no significant differences in abundance across salinities. For the mosquito species, we also estimated survival to pupation. Survival to pupation was significantly lower for O. caspius in freshwater, but was not statistically significantly different across salinities for A.␣multicolor. Species diversity was highest at the two lowest salinities tested and then dropped with increasing salinity. Evenness was not significantly different across salinities. Community similarity generally decreased with increasing salinity differences though dissimilarity was greatest when comparing freshwater to other salinities. Thus, regional diversity is likely increased when there is a range of salinities among pools.     

Molecular Analysis of the Sulfate Reducing and ArchaealCommunity in a Meromictic Soda Lake (Mono Lake, California) by Targeting 16S rRNA, mcrA, apsA, and dsrAB Genes

Sulfate reduction is the most important process involved in the mineralization of carbon in the anoxic bottom waters of Mono Lake, an alkaline, hypersaline, meromictic Lake in California. Another important biogeochemical process in Mono Lake is thought to be sulfate-dependent methane oxidation (SDMO). However little is known about what types of organisms are involved in these processes in Mono Lake. Therefore, the sulfate-reducing and archaeal microbial community in Mono Lake was analyzed by targeting 16S rRNA, methyl-coenzyme M reductase (mcrA), adenosine-5′-phosphosulfate (apsA), and dissimilatory sulfite reductase (dsrAB) genes to investigate the sulfate-reducing and archaeal community with depth. Most of the 16S rRNA gene sequences retrieved from the samples fell into the δ-subdivision of the Proteobacteria. Phylogenetic analyses suggested that the clones obtained represented sulfate-reducing bacteria, which are probably involved in the mineralization of carbon in Mono Lake, many of them belonging to a novel line of descent in the δ-Proteobacteria. Only 6% of the sequences retrieved from the samples affiliated to the domain Euryarchaeota but did not represent Archaea, which is considered to be responsible for SDMO [Orphan et al. 2001: Appl Environ Microbiol 67:1922–1934; Teske et al.: Appl Environ Microbiol 68:1994–2007]. On the basis of our results and thermodynamic arguments, we proposed that SDMO in hypersaline environments is presumably carried out by SRB alone. Polymerase chain reaction (PCR) amplifications of the mcrA-, apsA-, and dsrAB genes in Mono Lake samples were, in most cases, not successful. Only the PCR amplification of the apsA gene was partially successful. The amplification of these functional genes was not successful because there was either insufficient “target” DNA in the samples, or the microorganisms in Mono Lake have divergent functional genes.     

Selective feeding by baltic herring

Lake Lenore is a highly productive alkaline lake in the Lower Grand Coulee, Central Washington. Leakage of freshwater from nearby irrigation projects decreased the lake salinity from about 17 to 1.5 g/l during 1950–75. The initial bottom fauna with mass occurrence of single halobiontic chironomid species was supplanted by more diverse communities at all depths. The species richness increased by 2–8 times. The greatest addition of new species was found in the salinity range of 2–3 g/l. Concurrent increases in abundance and biomass suggest that the carrying capacity of the lake at higher salinities was not fully utilized by the benthos. A comparison with other eutrophic saline lakes indicates that benthic species composition and quantity is correlated with total salinity and ion composition.     

NITZSCHIA OVALIS (BACILLARIOPHYCEAE) MONO LAKE STRAIN ACCUMULATES 1,4/2,5 CYCLOHEXANETETROL IN RESPONSE TO INCREASED SALINITY1

The growth of microalgae in hypersaline conditions requires that cells accumulate osmoprotectants. In many instances, these are polyols. We isolated the diatom Nitzschia ovalis H. J. Arn. from the saline and alkaline water body Mono Lake (CA, USA). This isolate can grow in salinities ranging from 5 to 120 parts per thousand (ppt) of salt but normally at 90 ppt salinity. In this report, we identified the major polyol osmoprotectant as 1,4/2,5 cyclohexanetetrol by electron ionization‐mass spectrometry (EI–MS), ¹H, ¹³C nuclear magnetic resonance spectroscopy (NMR), and infrared (IR) and showed an increase in cellular concentration in response to rising salinity. This increase in the cyclitol concentration was evaluated by gas chromatography of the derived tetraacetylated cyclohexanetetrol obtaining an average of 0.7 fmol · cell^?1 at 5 ppt and rising to 22.5 fmol · cell^?1 at 120 ppt. The 1,4/2,5 cyclohexanetetrol was also detected in the red alga Porphyridium purpureum. Analysis of the free amino acid content in N. ovalis cultures exposed to changes in salinity showed that proline and lysine also accumulate with increased salinity, but the cellular concentration of these amino acids is about 10‐fold lower than the concentration of 1,4/2,5 cyclohexanetetrol. The comparison of amino acid concentration per cell with cyclitol suggests that this polyol is important in compensating the cellular osmotic pressure due to increased salinity, but other physiological functions could also be considered.