Hydrology of Lake Atnsjøen and River Atna

The hydrological investigations in the Atna basin started in 1917 with discharge measurements at the outlet of Lake Atnsjøen. Ice data are available from the lake since 1950 and water temperature data from 1980. With the start of the FORSKREF program in 1986 the number of hydrological stations increased substantially and included also snow measurements. During the FORSKREF period 1986–98 the annual mean discharge was 8% lower than during the normal period 1931–60. The annual flood is highest during the snow melt in spring, an exception was 1987 when a rain flood in October was the annual high. The spring flood in early June 1995 was the largest ever recorded and had a 100–200 year return period. The water temperature is low above tree-line at 1000 m a. s. l., rarely exceeding 7–8?°C. The temperature increases gradually with decreasing altitude and is periodically above 15?°C at 400 m a. s. l. The Atna Lake is an important heat source for the river downstream. In this lake, wind mixing normally prevents the development of a strong summer thermocline. The lake usually freezes over in late November and has a stable ice cover of 50–90 cm thickness in early spring. The ice normally breaks up in late May; thus the mean ice-covered period is 6 months. The ice cover on the river is less stable and rather large ice-runs are common on the lower part of River Atna. The snow measurements are carried out in April in a small subbasin between 800 and 1200 m a. s. l. The snow cover is unevenly distributed due to strong wind transport, but the pattern is more or less the same from one season to another. The mean snow accumulation in this basin is equivalent to 205 mm of precipitation.     

Spring time production of bottom ice algae in the landfast sea ice zone at Barrow, Alaska

The primary production of bottom ice algae is an important food source for sympagic, pelagic and benthic organisms in the Arctic Ocean as well as Antarctic Ocean. Using ¹³C-¹⁵N isotope tracers, the recent ice algal production at Barrow during the spring season was lower in 2003 than three decades ago, although the maximum chlorophyll-a concentration for the bottom ice algae was similar to the values from previous studies. Estimated recent new and total production rates of the ice algae were 0.8 g C m^- 2 yr^- 1 and 2.0 g C m^- 2 yr^- 1 respectively, while the rates of water column phytoplankton were 0.2 g C m^- 2 yr^- 1 and 0.7 g C m^- 2 yr^- 1 for the spring sampling period in 2003. The ice algae contributed 74% of the pelagic primary production under the landfast sea ice at Barrow before the phytoplankton spring bloom. At the end of the season in 2003, a high carbon allocation of lipids in the ice algae was found. Three possible explanations- nutrient depletion, increasing light, and/or changes in species composition- were suggested for the high carbon incorporation into lipids. This high lipid synthesis of the bottom ice algae might be significant to zooplankton and benthic fauna grazers because lipids are the most energy dense biomolecules.     

Ecosystem response to earlier ice break‐up date: Climate‐driven changes to water temperature,lake‐habitat‐specific production,and trout habitat and resource use

Climate warming has yielded earlier ice break‐up dates in recent decades for lakes leading to water temperature increases, altered habitat, and both increases and decreases to ecosystem productivity. Within lakes, the effect of climate warming on secondary production in littoral and pelagic habitats remains unclear. The intersection of changing habitat productivity and warming water temperatures on salmonids is important for understanding how climate warming will impact mountain ecosystems. We develop and test a conceptual model that expresses how earlier ice break‐up dates influence within lake habitat production, water temperatures and the habitat utilized by, resources obtained and behavior of salmonids in a mountain lake. We measured zoobenthic and zooplankton production from the littoral and pelagic habitats, thermal conditions, and the habitat use, resource use, and fitness of Brook Trout (Salvelinus fontinalis). We show that earlier ice break‐up conditions created a "resource‐rich" littoral–benthic habitat with increases in zoobenthic production compared to the pelagic habitat which decreased in zooplankton production. Despite the increases in littoral–benthic food resources, trout did not utilize littoral habitat or zoobenthic resources due to longer durations of warm water temperatures in the littoral zone. In addition, 87% of their resources were supported by the pelagic habitat during periods with earlier ice break‐up when pelagic resources were least abundant. The decreased reliance on littoral–benthic resources during earlier ice break‐up caused reduced fitness (mean reduction of 12 g) to trout. Our data show that changes to ice break‐up drive multi‐directional results for resource production within lake habitats and increase the duration of warmer water temperatures in food‐rich littoral habitats. The increased duration of warmer littoral water temperatures reduces the use of energetically efficient habitats culminating in decreased trout fitness.     

The inshore marine ecosystem off the Vestfold Hills,Antarctica

The planktonic, ice/water interface, and benthic communities at three sites off the coast of the Vestfold Hills, Antarctica, were examined over a complete year.The planktonic flora and fauna were composed predominantly of oceanic species with diatoms and copepods the numerically dominant groups. Primary production was largely restricted to the summer months except for epontic algae which developed in spring. The zooplankton exhibited a similar seasonal cycle but lagged some months behind that of the phytoplankton.The ice/water interface (epontic) fauna consisted of species from the plankton and benthos. Copepods were major contributors; however, two amphipod species dominated. Seasonality of the fauna in this habitat was determined by ice formation and breakout, and development of ice algae.Each of the benthic substrates supported a characteristic macrofaunal assemblage, although infaunal amphipods and tanaids were similar at each site. Infauna exhibited a distinct seasonal cycle related to that of the primary producers whereas macrofauna showed no seasonal changes in abundance.Species composition of each community in this coastal antarctic region was comparable with that of similar habitats in other antarctic coastal areas, supporting the circumpolarity of antarctic marine communities.     

Overwintering of Invertebrates in a Shallow Northern Swedish Lake

During two winters, the benthic invertebrates in a shallow northern Swedish lake (66 °N) were studied. The lake is ice-covered for about 200 days a year. Live Lamellibranchiata, Oligochaeta. Hydracarina, Isopoda, Heteroptera, Cole optera and Diptera were found in samples of ice and of frozen lake-bottom sediments. During the winter in which the bottom froze deepest, a 70–90% decrease in numbers of live Chironomidae larvae was recorded between the autumn and following early spring.     

Reproductive adaptations of Antarctic benthic invertebrates

The majority of Antarctic benthic invertebrates so far studied do not produce pelagic larvae, but develop non-pelagically by means of egg capsules, brooding or viviparity. The predominance of protected development in the Antarctic benthos is primarily due to the short period of summer phytoplankton abundance and the low sea temperature. Such conditions make it difficult for a larva to complete pelagic development before food becomes scarce in the surface waters. Prosobranch gastropods illustrate some important aspects of Antarctic benthic invertebrate reproduction. Species which develop non-pelagically have an aseasonal or prolonged spawning period. They produce a small number of large yolky eggs which remain in the benthos and develop slowly, giving rise to large, fully competent juveniles. Conversely, one species with free development has a short, synchronous spawning period during early summer, producing larvae which can benefit from the phytoplankton bloom. Protected development by means of brooding will limit dispersion, but transport on floating algae and by anchor ice may partially compensate for this in the Antarctic.     

Measurements of Seasonal Rates and Annual Budgets of Organic Carbon Fluxes in an Antarctic Coastal Environment at Signy Island, South Orkney Islands, Suggest a Broad Balance between Production and Decomposition

We report here the first comprehensive seasonal study of benthic microbial activity in an Antarctic coastal environment. Measurements were made from December 1990 to February 1992 of oxygen uptake and sulfate reduction by inshore coastal sediments at Signy Island, South Orkney Islands, Antarctica. From these measurements the rate of benthic mineralization of organic matter was calculated. In addition, both the deposition rate of organic matter to the bottom sediment and the organic carbon content of the bottom sediment were measured during the same period. Organic matter input to the sediment was small under winter ice cover, and the benthic respiratory activity and the organic content of the surface sediment declined during this period as available organic matter was depleted. On an annual basis, about 32% of benthic organic matter mineralization was anoxic, but the proportion of anoxic compared with oxic mineralization increased during the winter as organic matter was increasingly buried by the amphipod infauna. Fresh organic input occurred as the sea ice melted and ice algae biomass sedimented onto the bottom, and input was sustained during the spring after ice breakup by continued primary production in the water column. The benthic respiratory rate and benthic organic matter content correspondingly increased towards the end of winter with the input of this fresh organic matter. The rates of oxygen uptake during the southern summer (80 to 90 mmol of O₂ m^-2 day^-1) were as high as those reported for other sediments at much higher environmental temperatures, and the annual mineralization of organic matter was equally high (12 mol of C m^-2 year^-1). Seasonal variations of benthic activity in this antarctic coastal sediment were regulated by the input and availability of organic matter and not by seasonal water temperature, which was relatively constant at between -1.8 and 0.5°C. We conclude that despite the low environmental temperature, organic matter degradation broadly balanced organic matter production, although there may be significant interrannual variations in the sources of the organic matter inputs.     

Disturbance, colonization and development of Antarctic benthic communities

A decade has yielded much progress in understanding polar disturbance and community recovery-mainly through quantifying ice scour rates, other disturbance levels, larval abundance and diversity, colonization rates and response of benthos to predicted climate change. The continental shelf around Antarctica is clearly subject to massive disturbance, but remarkably across so many scales. In summer, millions of icebergs from sizes smaller than cars to larger than countries ground out and gouge the sea floor and crush the benthic communities there, while the highest wind speeds create the highest waves to pound the coast. In winter, the calm associated with the sea surface freezing creates the clearest marine water in the world. But in winter, an ice foot encases coastal life and anchor ice rips benthos from the sea floor. Over tens and hundreds of thousands of years, glaciations have done the same on continental scales-ice sheets have bulldozed the seabed and the zoobenthos to edge of shelves. We detail and rank modern disturbance levels (from most to least): ice; asteroid impacts; sediment instability; wind/wave action; pollution; UV irradiation; volcanism; trawling; non-indigenous species; freshwater inundation; and temperature stress. Benthic organisms have had to recolonize local scourings and continental shelves repeatedly, yet a decade of studies have demonstrated that they have (compared with lower latitudes) slow tempos of reproduction, colonization and growth. Despite massive disturbance levels and slow recolonization potential, the Antarctic shelf has a much richer fauna than would be expected for its area. Now, West Antarctica is among the fastest warming regions and its organisms face new rapid changes. In the next century, temperature stress and non-indigenous species will drastically rise to become dominant disturbances to the Antarctic life. Here, we describe the potential for benthic organisms to respond to disturbance, focusing particularly on what we know now that we did not a decade ago.     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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Annual cycle of phytoplankton in Ace Lake,an ice covered,saline meromictic lake

The annual cycle of phytoplankton in saline, meromictic Ace Lake (68°2S.4S, 78°11.1E) in the Vestfold Hills, Antarctica, was studied from January, 1979 to January 1980. Ace Lake has permanent gradients of temperature, salinity, dissolved oxygen, and hydrogen sulphide, and is ice covered with up to 2 m of ice for 10–12 months each year. The phytoplankton community had low diversity, consisting of only four species, all flagellates — a prasinophyte Pyramimonas gelidicola McFadden et al., a cryptophyte of the genus Cryptomonas; an unidentified colourless microflagellate, and an unarmoured dinoflagellate. These were restricted to the oxic zone of the lake from the surface to 10 m.The phytoplankton had a cycle of seven months of active growth over spring and summer. Low numbers of cells survived in the water column over winter. Spring growth was initiated below the ice by increased light penetration through the ice into the lake, enhanced at the time by the removal of surface snow which accumulated on the ice over winter. Peak phytoplankton biomass production was by the shade adapted P. gelidicola and occurred at the interface of the oxic and anoxic zones where substantial available nitrogen as ammonia is found.The three dominant phytoplankton species displayed distinct vertical stratification over the oxic zone. This stratification was not static and developed over spring as the flagellates migrated to preferred light climate zones. Mean cell volume of two of the flagellates varied significantly over the year. Minimum volumes were recorded in winter and volume increased progressively over spring to reach maximum mean cell volume in summer. Mean cell volume was positively correlated with light intensity (maximum ambient PAR at the respective depth for date of sample). Low cell volume in winter may be related to winter utilization of carbohydrate reserves by slow respiration, and may represent a survival mechanism.     

Effects of temperature and sediment properties on benthic CO2 production in an oligotrophic boreal lake

1. Temperature and many other physical and chemical factors affecting CO₂ production in lake sediments vary significantly both seasonally and spatially. The effects of temperature and sediment properties on benthic CO₂ production were studied in in situ and in vitro experiments in the boreal oligotrophic Lake Pääjärvi, southern Finland. 2. In in situ experiments, temperature of the water overlying the shallow littoral sediment varied seasonally between 0.5 and 15.7 °C, but in deep water (≥20 m) the range was only 1.1–6.6 °C. The same exponential model (r² = 0.70) described the temperature dependence at 1.2, 10 and 20 m depths. At 2.5 and 5 m depths, however, the slopes of the two regression models (r² = 0.94) were identical but the intercept values were different. Sediment properties (wet, dry, mineral and organic mass) varied seasonally and with depth, but they did not explain a significantly larger proportion of variation in the CO₂ output rate than temperature. 3. In in vitro experiments, there was a clear and uniform exponential dependence of CO₂ production on temperature, with a 2.7‐fold increase per 10 °C temperature rise. The temperature response (slope of regression) was always the same, but the basic value of CO₂ production (intercept) varied, indicating that other factors also contributed to the benthic CO₂ output rate. 4. The annual CO₂ production of the sediment in Lake Pääjärvi averaged 62 g CO₂ m^?2, the shallow littoral at 0–3 m depth releasing 114 g CO₂ m^?2 and deep profundal (>15 m) 30 g CO₂ m^?2. On the whole lake basis, the shallow littoral at 0–3 m depth accounted for 53% and the sediment area in contact with the summer epilimnion (down to a depth c. 10 m) 75% of the estimated total annual CO₂ output of the lake sediment, respectively. Of the annual production, 83% was released during the spring and summer. 5. Using the temperature‐CO₂ production equations and climate change scenarios we estimated that climatic warming might increase littoral benthic CO₂ production in summer by nearly 30% from the period 1961–90 to the period 2071–2100.     

Effects of spates on interstitial assemblages of the Rhône River. Importance of spatial heterogeneity

The dynamics of interstitial assemblages, after a spate and during low discharge, was studied in a regulated channel (Miribel Canal) of the Upper Rhône River, France. Using a Bou-Rouch pump, three stations were samples: 1) Station IIA, a site fed by superficial water infiltrations, 2) Station IIC, a site fed by riparian phreatic water, and 3) Station IA, a site fed by both surface and phreatic waters. The spate greatly influenced the interstitial assemblages, their dynamics were different according to the hydrology of the given site. At Station IIA, the spate had a wash-out effect on the assemblages (reduction in abundance and diversity), whereas during low discharge the interstitial layer received a continuous influx of epigean organisms (benthic and limnophilous). At Stations IIC and IA, the spate introduced numerous limnophilous and benthic invertebrates into the interstices, which function as an organismic trap. However, these sites appear to be more isolated from surface waters during low discharge. Stygobites decreased or disappeared after the spate. They appear highly sensitive to hydrologic perturbations in the surface waters.     

A three-dimensional general circulation model of the surface layers of Lake Baikal

A three-dimensional general circulation model has been developed to investigate mixing processes in Lake Baikal, Siberia. Emphasis is placed on the 4–5-month period when the lake is completely ice-covered, a time of particular importance to the re-population of the lake by diatoms. The model represents the top 250 m and includes a new mixing scheme developed specifically for the investigation of under-ice flows. The effects of spatial patterns of snow and ice transparency on circulation and temperature are investigated. In general, temperature profiles provide an indication of the extent and depth of mixing and are highly sensitive to the presence of snow and to the transparency of ice. Generated profiles agree well with in situ measurements, which are difficult to obtain during this period. The model is shown to be particularly successful in simulating mixing processes in Lake Baikal. The surface heat fluxes that are required for a model of this type were estimated using satellite data, which provide complete coverage of the lake within one image. An increase in albedo values of 20% has no significant impact on the development of the temperature profile. Finally, density driven currents generated in the model were investigated. The magnitudes of the model currents compared to observations suggest that the background flow under ice in the lake may be density driven.     

Long-term trends in water temperature and ice cover in the subalpine lake, Øvre Heimdalsvatn,and nearby lakes and rivers

Long-term data series of ice cover on lakes and river temperatures from the mountain areas of Norway are lacking. The present study analyses the last four decades of ice data from the subalpine lake, Øvre Heimdalsvatn, and water temperature data from its outlet river, Hinøgla. These data are compared to water temperature data from three neighbouring, quite different locations, the glacier-fed rivers Leirungsåi and Sjoa, and the alpine lake, Bessvatn. The study also examines the air temperature/river temperature relationships, and the air temperature/ice freeze-up and break-up dates. During the months of July, August and September, the water temperature in Hinøgla was well correlated to the air temperature, but the correlation was poor in the remaining months due to the ice cover and snow conditions. A significant temperature increase of 2–3°C has been observed in Hinøgla in the months August–October since 1984. There were only minor changes in the duration of the ice cover season during the last 40 years, but a delay of 9 days was found in the freeze-up date and a delay of 6 days in the break-up date, although the latter was not significant.     

Variation in spatial and temporal gradients in zooplankton spring development: the effect of climatic factors

1. We examined the temporal and spatial heterogeneity of zooplankton in lake surface waters during the spring of 3 years in Lake Washington, U.S.A., a large lake with a high production of sockeye salmon fry. 2. We show large within‐season and among‐year variation in the horizontal distribution of temperature, chlorophyll a concentration, and zooplankton in the lake. The main pattern, a delay in zooplankton population increase from the north‐ to the south‐end of the lake, recurred in each year and was persistent within each spring. 3. The delay is primarily caused by the development of a temperature gradient during spring warming, as cold mountain water enters the south end of the lake, while warm water enters the north end via a river draining a nearby lake. Climate factors, such as air temperature and precipitation during winter and spring, appear to influence the extent of the delay of zooplankton increase. 4. If the climate continues to warm, the temporal disconnection in zooplankton development between lake areas immediately influenced by cold river inflow and areas that are influenced by spring warming may increase in magnitude. Thus, the different areas of the lake may not contribute equally to fish production.     

The life cycle dynamics and production of zooplankton in Øvre Heimdalsvatn

The zooplankton in Øvre Heimdalsvatn was studied from May 1969 to Aug 1973 in order to obtain knowledge about their functioning and role in the lake ecosystem. The paper concentrates on the main species: Holopedium gibberum, Bosmina longispina, Cyclops scutifer. Heterocope saliens, Conochilus unicornis, Polyarthra vulgaris and Kellicottia longispina but Megacyclops gigus, Daphnia longispina and ciliated cells are also considered. The development of the populations was fairly synchronized and made it possible to determine life cycles, development times, growth, reproduction and production from field data. The total production of the multicellular zooplankton was 2–3 mg dry wt m⁻² yr⁻¹. The cladocerans dominated and their share of the total was 84–92% compared to the copepods 5–8% and the rotifers 3–8%. The ciliate production was not estimated, but their biomass exceeded that of the copepods and rotifers. The spring spate, June temperatures, food availability and invertebrate predation were found to be the most important regulatory factors for life cycle dynamics and production. Allochthonous organic matter was found to be a necessary food resource in addition to algae.     

SEASONAL PERIODICITY OF CHRYSOPHYCEAE AND SYNUROPHYCEAE IN A SMALL NEW ENGLAND LAKE: IMPLICATIONS FOR PALEOLIMNOLOGICAL RESEARCH1

The seasonal periodicity of taxa of Chrysophyceae and Synurophyceae from a small New England lake is described for the period September 1983 through June 1988. We found 51 taxa, including 29 that accounted for over 10% of the total in at least one collection. The taxa were fitted into one of five seasonal patterns. Patterns I and II represented taxa restricted to warm (pattern I) or cold (pattern II) months, respectively. Pattern HI represented organisms that began growth in the summer, persisted through autumn and disappeared with the onset of an ice cover. Pattern IV was an extension of pattern III, in which the taxon remained in the plankton throughout the winter and disappeared soon after ice out. Species without a clear seasonal pattern were grouped as pattern V. The seasonal periodicity of the flora, as examined with ordination analyses, was found to remain remarkedly similar during the 58–month study. Except for episodes of low pH during spring snow melt and unseasonally warm or cold weather, sample scores followed a fairly consistent pattern along the first and second primary axes. Water temperature, specific conductance, and pH were important variables that controlled changes in the species composition during the course of a given year. The flora was used to develop an inference model for water temperature. According to the analyses, the remains of a surface sediment sample represented a flora that grew primarily during the late autumn period at 7.6 ° C. Ways in which seasonal data could be utilized to improve paleolimnological inference work are discussed.     

The ecology of Lake Myvatn and the River Laxá: Variation in space and time

Ecological features of Lake Myvatn and the outflowing River Laxá show a wide range of spatial and temporal variations. The physical division of the lake into three main basins and the variation in chemical composition and temperature of the artesian springs feeding this shallow productive lake have large spatial effects. Variation in groundwater characteristics depends on percolation time and proximity to geothermal sources. Variation in precipitation is evened out by the porous volcanic soil and bedrock and the spring-water discharge is therefore very stable. A pulse of volcanic activity in 1975–1984 (the Krafla Fires) heated the groundwater entering the North Basin of the lake and changed its chemistry. Although much reduced, these effects have not disappeared yet, but overall the impact of the volcanic activity on the biota seemed minimal. Recycling of nutrients through internal loading is important and occurs on various time scales. In winter, when the lake is ice-covered, the topmost 5-cm layer of sediment pore water has a hundredfold concentration of nutrients relative to the overlying lake water. The nutrients are released during the ice-free period by sediment resuspension, diffusion, bioturbation and recycling. In spring, resuspension events sometimes lead to spikes in dissolved phosphorus and nitrogen, but there is little evidence of any major desorption of nutrients from suspended particles during such events later in the summer. In contrast to the stable groundwater, the biota show more or less regular fluctuations with no straightforward correlation with external signals. The most prominent fluctuations, those of the chironomid Tanytarsus gracilentus seem to be driven by interactions between the species and its sediment resources. Fluctuations in other invertebrates could be a consequence of the Tanytarsus cycles due to the large impact this species has on the benthic environment of this detritus-driven ecosystem. Temporal variation in epibenthic chironomids and Cladocera translates into variable production of vertebrate predators (Arctic charr, Salvelinus alpinus, and ducks), body condition and mortality of fish and sometimes into return rates of migrating adult ducks. The waterfowl show large temporal variation on a centennial scale, e.g., the invasion of the tufted duck (Aythya fuligula) which arrived by the end of the 19th century and has by now outnumbered other species. Fluctuations of Cyanobacteria (Anabaena) and the fish Gasterosteus aculeatus (three-spined stickleback) harmonize with the cycles in the benthic community. Palaeolimnological studies indicate that primary production in the South Basin became increasingly benthic as the lake depth was reduced by sedimentation (around 2 mm year^–1). Other trends include a decrease in Tanytarsus and Daphnia and an exponential increase in green algae (Cladophorales, Pediastrum) and associated organisms.     

Anchor ice and benthic disturbance in shallow Antarctic waters: interspecific variation in initiation and propagation of ice crystals

Sea ice typically forms at the ocean's surface, but given a source of supercooled water, an unusual form of ice--anchor ice--can grow on objects in the water column or at the seafloor. For several decades, ecologists have considered anchor ice to be an important agent of disturbance in the shallow-water benthic communities of McMurdo Sound, Antarctica, and potentially elsewhere in polar seas. Divers have documented anchor ice in the McMurdo communities, and its presence coincides with reduced abundance of the sponge Homaxinella balfourensis, which provides habitat for a diverse assemblage of benthic organisms. However, the mechanism of this disturbance has not been explored. Here we show interspecific differences in anchor-ice formation and propagation characteristics for Antarctic benthic organisms. The sponges H. balfourensis and Suberites caminatus show increased incidence of formation and accelerated spread of ice crystals compared to urchins and sea stars. Anchor ice also forms readily on sediments, from which it can grow and adhere to organisms. Our results are consistent with, and provide a potential first step toward, an explanation for disturbance patterns observed in shallow polar benthic communities. Interspecific differences in ice formation raise questions about how surface tissue characteristics such as surface area, rugosity, and mucus coating affect ice formation on invertebrates.