Food resource effects on diel movements and body size of cisco in north-temperate lakes

The movement patterns and body size of fishes are influenced by a host of physical and biological conditions, including temperature and oxygen, prey densities and foraging potential, growth optimization, and predation risk. Our objectives were to (1) investigate variability in vertical movement patterns of cisco (Coregonus artedi) in a variety of inland lakes using hydroacoustics, (2) explore the causal mechanisms influencing movements through the use of temperature/oxygen, foraging, growth, and predation risk models, and (3) examine factors that may contribute to variations in cisco body size by considering all available information. Our results show that cisco vertical movements vary substantially, with different populations performing normal diel vertical migrations (DVM), no DVM, and reverse DVM in lakes throughout Minnesota and northern Wisconsin, USA. Cisco populations with the smallest body size were found in lakes with lower zooplankton densities. These smaller fish showed movements to areas of highest foraging or growth potential during the day and night, despite moving out of preferred temperature and oxygen conditions and into areas of highest predation risk. In lakes with higher zooplankton densities, cisco grew larger and had movements more consistent with behavioral thermoregulation and predator avoidance, while remaining in areas with less than maximum foraging and growth potential. Furthermore, the composition of potential prey items present in each lake was also important. Cisco that performed reverse DVM consumed mostly copepods and cladocerans, while cisco that exhibited normal DVM or no migration consumed proportionally more macro-zooplankton species. Overall, our results show previously undocumented variation in migration patterns of a fish species, the mechanisms underlying those movements, and the potential impact on their growth potential.     

Behavioural thermoregulation by subyearling fall (autumn) Chinook salmon Oncorhynchus tshawytscha in a reservoir

This study investigated behavioural thermoregulation by subyearling fall (autumn) Chinook salmon Oncorhynchus tshawytscha in a reservoir on the Snake River, Washington, U.S.A. During the summer, temperatures in the reservoir varied from 23° C on the surface to 11° C at 14 m depth. Subyearlings implanted with temperature-sensing radio transmitters were released at the surface at temperatures >20° C during three blocks of time in summer 2004. Vertical profiles were taken to measure temperature and depth use as the fish moved downstream over an average of 5·6–7·2 h and 6·0–13·8 km. The majority of the subyearlings maintained average body temperatures that differed from average vertical profile temperatures during most of the time they were tracked. The mean proportion of the time subyearlings tracked within the 16–20° C temperature range was larger than the proportion of time this range was available, which confirmed temperature selection opposed to random use. The subyearlings selected a depth and temperature combination that allowed them to increase their exposure to temperatures of 16–20° C when temperatures <16 and >20° C were available at lower and higher positions in the water column. A portion of the subyearlings that selected a temperature c. 17·0° C during the day, moved into warmer water at night coincident with an increase in downstream movement rate. Though subyearlings used temperatures outside of the 16–20° C range part of the time, behavioural thermoregulation probably reduced the effects of intermittent exposure to suboptimal temperatures. By doing so, it might enhance growth opportunity and life-history diversity in the population of subyearlings studied.     

The highest kingdom of Anolis: Thermal biology of the Andean lizard Anolis heterodermus (Squamata: Dactyloidae) over an elevational gradient in the Eastern Cordillera of Colombia

Vertebrate ectotherms may deal with changes of environmental temperatures by behavioral and/or physiological mechanisms. Reptiles inhabiting tropical highlands face extreme fluctuating daily temperatures, and extreme values and intervals of fluctuations vary with altitude. Anolis heterodermus occurs between 1800 m to 3750 m elevation in the tropical Andes, and is the Anolis species found at the highest altitude known. We evaluated which strategies populations from elevations of 2200 m, 2650 m and 3400 m use to cope with environmental temperatures. We measured body, preferred, critical maximum and minimum temperatures, and sprint speed at different body temperatures of individuals, as well as operative temperatures. Anolis heterodermus exhibits behavioral adjustments in response to changes in environmental temperatures across altitudes. Likewise, physiological traits exhibit intrapopulation variations, but they are similar among populations, tended to the “static” side of the evolution of thermal traits spectrum. The thermoregulatory behavioral strategy in this species is extremely plastic, and lizards adjust even to fluctuating environmental conditions from day to day. Unlike other Anolis species, at low thermal quality of the habitat, lizards are thermoconformers, particularly at the highest altitudes, where cloudy days can intensify this strategy even more. Our study reveals that the pattern of strategies for dealing with thermal ambient variations and their relation to extinction risks in the tropics that are caused by global warming is perhaps more complex for lizards than previously thought.     

Impact of global warming at the range margins: phenotypic plasticity and behavioral thermoregulation will buffer an endemic amphibian

When dispersal is not an option to evade warming temperatures, compensation through behavior, plasticity, or evolutionary adaptation is essential to prevent extinction. In this work, we evaluated whether there is physiological plasticity in the thermal performance curve (TPC) of maximum jumping speed in individuals acclimated to current and projected temperatures and whether there is an opportunity for behavioral thermoregulation in the desert landscape where inhabits the northernmost population of the endemic frog Pleurodema thaul. Our results indicate that individuals acclimated to 20°C and 25°C increased the breath of their TPCs by shifting their upper limits with respect to when they were acclimated at 10°C. In addition, even when dispersal is not possible for this population, the landscape is heterogeneous enough to offer opportunities for behavioral thermoregulation. In particular, under current climatic conditions, behavioral thermoregulation is not compulsory as available operative temperatures are encompassed within the population TPC limits. However, for severe projected temperatures under climate change, behavioral thermoregulation will be required in the sunny patches. In overall, our results suggest that this population of Pleurodema thaul will be able to endure the worst projected scenario of climate warming as it has not only the physiological capacities but also the environmental opportunities to regulate its body temperature behaviorally.     

Browning affects pelagic productivity in northern lakes by surface water warming and carbon fertilization

Global change impacts important environmental drivers for pelagic gross primary production (GPP) in northern lakes, such as temperature, light, nutrient, and inorganic carbon availability. Separate and/or synergistic impacts of these environmental drivers on pelagic GPP remain largely unresolved. Here, we assess key drivers of pelagic GPP by combining detailed depth profiles of summer pelagic GPP with environmental and climatic data across 45 small and shallow lakes across northern Sweden (20 boreal, 6 subarctic, and 19 arctic lakes). We found that across lakes summer pelagic GPP was strongest associated with lake water temperatures, lake carbon dioxide (CO₂) concentrations impacted by lake water pH, and further moderated by dissolved organic carbon (DOC) concentrations influencing light and nutrient conditions. We further used this dataset to assess the extent of additional DOC-induced warming of epilimnia (here named internal warming), which was especially pronounced in shallow lakes (decreasing 0.96°C for every decreasing m in average lake depth) and increased with higher concentrations of DOC. Additionally, the total pools and relative proportion of dissolved inorganic carbon and DOC, further influenced pelagic GPP with drivers differing slightly among the boreal, subarctic and Arctic biomes. Our study provides novel insights in that global change affects pelagic GPP in northern lakes not only by modifying the organic carbon cycle and light and nutrient conditions, but also through modifications of inorganic carbon supply and temperature. Considering the large-scale impacts and similarities of global warming, browning and recovery from acidification of lakes at higher latitudes throughout the northern hemisphere, these changes are likely to operate on a global scale.     

Modification of the diel vertical migration of Bythotrephes longimanus by the cold-water planktivore, Coregonus artedi

1. The weak diel vertical migration observed in the large cladoceran Bythotrephes longimanus seems contradictory to the predator‐avoidance hypothesis that predicts large zooplankton should have long migration amplitudes. However, cold‐water planktivores, especially Coregonus spp., are a main source of mortality for Bythotrephes and hence a deeper migration would result in a greater overlap with these hypolimnetic planktivores. We hypothesized that Coregonus artedi (cisco) modifies the normal vertical migration pattern of Bythotrephes, such that the latter stays higher in the water column during the day and thus migrates less extremely at night. 2. The vertical distribution of Bythotrephes during the day was determined from single visits to six lakes in Ontario, Canada, all of which contain warm‐water, epilimnetic planktivores but differing in whether they contain cisco. One lake of each fish type was sampled day and night every 2–3 weeks over the ice‐free season to examine daytime depths and migration amplitude. 3. The vertical migration of Bythotrephes differed in the presence and absence of cisco. In the lakes with cisco, there were significantly fewer Bythotrephes in the hypolimnion and they were higher in the water column during the day. Migration amplitude was smaller in the cisco than in the non‐cisco lake. These observations were not attributable to differences in physical factors, and, although not conclusively attributable to cisco, are consistent with an effect of cisco. 4. We suggest that diurnal depth selection by Bythotrephes in lakes containing cisco is a trade‐off between the risk of predation by warm‐ versus cold‐water predators, balanced by the benefits of increased temperature and feeding rates near the surface. Even in lakes without cisco, however, the vertical migration of Bythotrephes was less than expected, suggesting that diurnal depth selection is a balance between the risk from warm‐water planktivores and access to sufficient light to feed effectively.     

The vertical distribution of fish in the open water area of a deep temperate mesotrophic lake assessed by hydroacoustics and midwater trawling

The fish stock of a deep temperate, mesotrophic lake was sampled at different depths using a fixed‐frame fry trawl, during two nights in mid‐September 2009. Additionally, horizontal and vertical hydroacoustics were used simultaneously to evaluate fish abundance and biomass estimates obtained by the trawl. Roach Rutilus rutilus and smelt Osmerus eperlanus were the dominant species of young‐of‐the‐year (YOY) fish in the trawl catches from the surface layers (0–9 m). Bleak Alburnus alburnus dominated the catch of older fish in the upper part of the surface profile (0–6 m). Around the thermocline (9–13 m) smelt dominated the catches of both the YOY and older fish. Beneath the thermocline (13–36 m) vendace Coregonus albula dominated the catch of YOY fish, and smelt was the only species of older fish in the trawl catches. Species composition, abundance and biomass of the YOY and older fish were heterogeneous throughout the depth profiles of the lake, but only abundance differed significantly between the layers. The hydroacoustics gave relatively similar estimates of abundance and biomass to those obtained by the trawl in all the depths sampled. Our results indicate that there is a clear separation of small fish of different species along the vertical profile of a deep temperate lake during the night, and an unequal vertical distribution of fish abundance and biomass. The similarity of the trawl and hydroacoustics estimates of abundances and biomass indicated that the trawl sampling did not cause important avoidance reactions of small fish during the night in this deep temperate lake (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Genetic differentiation of coregonid fishes in pechora river

A population–genetic study of five coregonid fish species has been carried out using 30 enzyme loci. The species under study included whitefish, a natural hybrid of vendace and least cisco, inconnu, peled, and Arctic cisco. The investigation revealed a low intraspecific genetic differentiation of the abovementioned species from Lower Pechora River and Usa River, which is the main tributary in the Pechora River basin, where coregonid fishes migrate to spawn. Presumably, whitefish, vendace, and inconnu inhabiting the main channels of Lower Pechora and Usa rivers are represented here by introgressive populations of species formed by a mix of discrete evolutionary lineages originating from geographically separated Late Quaternary refugia. Some lineages of whitefish, vendace, and inconnu from the West Siberian periglacial lake came to Eastern Europe together with the last migration wave of Siberian species to the west, including peled, broad whitefish, and Arctic cisco, during the Middle Weichselian glaciation (60000–50000 years ago), while in the Eastern European Periglacial refugium (Lake Komi), local lineages of these species had survived. The periglacial Lake Komi has probably been located on the Pechora Plain since early Weichselian glaciation (90000–80000 years ago), and the studied whitefish specimens from Upper Pechora River are probably the purest descendants of the periglacial race of whitefish from Lake Komi. This assumption is supported by the data on the high level of genetic differentiation between the whitefish of Upper Pechora and Lower Pechora (D _N = 0.005), as well as by the pattern of allele distribution in some polymorphic loci. In our opinion, the ecologic and genetic differentiation of vendace in the Pechora River basin is also connected with the interaction between two evolutionary lineages that originated from independent periglacial refugia.     

Latitudinal variation in thermal ecology of North American ratsnakes and its implications for the effect of climate warming on snakes

Behavioral thermoregulation is expected to be critical in determining the capacity of reptiles to respond to climate warming and how that response will vary with latitude. We used radio-telemetry to compare behavioral thermoregulation among ratsnake (Elaphe obsoleta) populations in Texas, Illinois, and Ontario, a latitudinal distance of >1500 km. Despite numerous specific differences among populations, overall the thermal ecology was surprisingly similar during the months that snakes in all three populations were active. Preferred temperatures varied only slightly across the snakes’ range, the extent of thermoregulation was similar, and by varying when during the day and season they thermoregulated, snakes in all three populations realized body temperatures within their preferred temperature range 15–20% of the time. The ability to use fine-scale behavioral thermoregulation (i.e., selective use of habitats and microclimates) to a similar extent and achieve similar outcomes across such a wide latitudinal and climatic gradient is made possible by large-scale differences in timing of activity (ratsnakes in Texas switch to nocturnal activity during summer, whereas in Illinois and Ontario activity is exclusively diurnal and hibernation lasts 5–7 months). Modeling indicated that a 3 °C increase in ambient temperature will generally improve thermal conditions for all three populations. Our empirical analyses suggest that the snakes’ ability to respond to climate warming will be determined more by their capacity to adjust when they are active than by changes in the extent of fine-scale behavioral thermoregulation. The ability to adjust timing of activity appears to make many snakes fundamentally different from lizards. As such, the consequences of climate warming may be very different for these two groups of reptiles.     

Comparing climate change and species invasions as drivers of coldwater fish population extirpations

Species are influenced by multiple environmental stressors acting simultaneously. Our objective was to compare the expected effects of climate change and invasion of non-indigenous rainbow smelt (Osmerus mordax) on cisco (Coregonus artedii) population extirpations at a regional level. We assembled a database of over 13,000 lakes in Wisconsin, USA, summarising fish occurrence, lake morphology, water chemistry, and climate. We used A1, A2, and B1 scenarios from the Intergovernmental Panel on Climate Change (IPCC) of future temperature conditions for 15 general circulation models in 2046-2065 and 2081-2100 totalling 78 projections. Logistic regression indicated that cisco tended to occur in cooler, larger, and deeper lakes. Depending upon the amount of warming, 25-70% of cisco populations are predicted to be extirpated by 2100. In addition, cisco are influenced by the invasion of rainbow smelt, which prey on young cisco. Projecting current estimates of rainbow smelt spread and impact into the future will result in the extirpation of about 1% of cisco populations by 2100 in Wisconsin. Overall, the effect of climate change is expected to overshadow that of species invasion as a driver of coldwater fish population extirpations. Our results highlight the potentially dominant role of climate change as a driver of biotic change.     

The size of a vendace, Coregonus albula L., stock in a deep lake basin monitored by hydroacoustic methods

An acoustic estimate of the number and biomass of a vendace stock (age >2 + years) in a deep basin of Lake Karjalan Pyhäjärvi was made in August 1985. The acoustic data were collected at night during the summer stagnation. The vendace were on the lake bottom during the day and rose by midnight to the hypolimnion; they did not rise to the thermocline or water layers above it. In the research area the mean number of vendace was 1900 fish ha ⁻¹ and the mean biomass was 76 kg ha ⁻¹. Total vendace biomass was 151 in the whole 200-ha research area.     

Ecophysiological responses to temperature of the "killer shrimp" Dikerogammarus villosus: is the invader really stronger than the native Gammarus pulex?

With global climate changes, biological invasions are considered to be one of the main causes of the decline of freshwater biodiversity. In this context, predicted increases in global temperature may alter the geographical distributions of native and invasive species. The purpose of our study was to examine the metabolic, behavioral and physiological responses to short-term temperature acclimation of two widely distributed species (the most successful European invader, Dikerogammarus villosus, and its main victim, Gammarus pulex), in order to estimate the potential effect of global warming on its invasion of freshwater ecosystems. Our results show that D. villosus is more vulnerable to high temperatures than G. pulex. The native species seems to be best adapted to intermediate temperatures (10-20°C) with a possibility of adjustment to "extreme" temperatures (5-27°C), whereas the "killer shrimp" D. villosus seems best adapted to lower temperatures (5-10°C) with a limited possibility of adjustment above 20°C. In the light of our results, global warming is likely to be less favorable to the invasive species. However, D. villosus showed reduced metabolic and activity rates, associated with higher glycogen content. This adaptive strategy was interpreted as having functional advantages, allowing D. villosus to successfully invade harsh and/or unpredictable biotopes. In addition, our results show that glycogen stores may be used as a powerful indicator of the optimal thermal window for aquatic ectotherms.     

High temperature acclimation of C4 photosynthesis is linked to changes in photosynthetic biochemistry

With average global temperatures predicted to increase over the next century, it is important to understand the extent and mechanisms of C4 photosynthetic acclimation to modest increases in growth temperature. To this end, we compared the photosynthetic responses of two C4 grasses (Panicum coloratum and Cenchrus ciliaris) and one C4 dicot (Flaveria bidentis) to growth at moderate (25/20 degrees C, day/night) or high (35/30 degrees C, day/night) temperatures. In all three C4 species, CO2 assimilation rates (A) underwent significant thermal acclimation, such that when compared at growth temperatures, A increased less than what would be expected given the strong response of A to short-term changes in leaf temperature. Thermal photosynthetic acclimation was further manifested by an increase in the temperature optima of A, and a decrease in leaf nitrogen content and leaf mass per area in the high- relative to the moderate-temperature-grown plants. Reduced photosynthetic capacity at the higher growth temperature was underpinned by selective changes in photosynthetic components. Plants grown at the higher temperature had lower amounts of ribulose-1,5-bisphosphate carboxylase/oxygenase and cytochrome f and activity of carbonic anhydrase. The activities of photosystem II (PSII) and phosphoenolpyruvate carboxylase were not affected by growth temperature. Chlorophyll fluorescence measurements of F. bidentis showed a corresponding decrease in the quantum yield of PSII (phi(PSII)) and an increase in non-photochemical quenching (phi(NPQ)). It is concluded that through these biochemical changes, C4 plants maintain the balance between the various photosynthetic components at each growth temperature, despite the differing temperature dependence of each process. As such, at higher temperatures photosynthetic nitrogen use efficiency increases more than A. Our results suggest C4 plants will show only modest changes in photosynthetic rates in response to changes in growth temperature, such as those expected within or between seasons, or the warming anticipated as a result of global climate change.     

Modelling lake cyanobacterial blooms: Disentangling the climate‐driven impacts of changing mixed depth and water temperature

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Application of a numerical model to predict impacts of climate change on water temperatures in two deep, oligotrophic lakes in New Zealand

We applied a numerical hydrodynamic model (DYRESM) to two large, deep New Zealand lakes that are characterised by deep thermoclines and high wind forcing, to assess their sensitivity to changes in climate. Modifications to standard model parameters were necessary for the successful application of DYRESM. Predictions from downscaled global circulation models suggest an increase in mean air temperature, rainfall, and wind speeds. Modelling the hydrodynamics of the lakes suggests that increasing air temperatures would offset the cooling influences of increased rainfall and river flows, resulting in warmer overall lake temperatures, and an earlier, longer, and shallower thermal stratification. These physical changes could affect phytoplankton production as their light limitation would decrease in duration and intensity. However, deeper mixing caused by increases in wind speed would negate this reduction of thermocline depth. While warmer air temperatures appear to be the dominant driver of changes in thermal structure, changes in other meteorological factors, especially wind speed, are important in predicting future hydrodynamics. Compared to large, deep lakes in the Northern Hemisphere, the predicted warming rates in Lakes Wanaka and Wakatipu are slower, due partly to a lower predicted rate of atmospheric warming and the absence of winter ice cover in these lakes.     

青藏高原多年冻土区不同水分条件的高寒草甸根系功能性状对增温的响应

在青藏高原多年冻土广泛分布的风火山地区,选择小嵩草（Kobresia pygmea）草甸和藏嵩草（Kobresia tibetica）沼泽化草甸为研究对象,采用开顶增温室（Open top chambers, OTCs）模拟气候变暖,探讨模拟增温对土壤水分差异的两种草甸地下生物量及根系功能性状的影响。结果显示,（1）增温显著增加小嵩草草甸0—20 cm根系生物量,主要是由于表层（0—10 cm）根系生物量显著增加,而对藏嵩草沼泽化草甸根系生物量无影响。（2）增温显著增加了小嵩草草甸根组织密度,同时提高了藏嵩草沼泽化草甸10—20 cm的比根长和比根面积（3）增温降低了小嵩草草甸的根系碳含量及10—20 cm根系氮含量,增加了藏嵩草沼泽化草甸的碳含量及10—20 cm根系氮含量,显著提高了小嵩草草甸和藏嵩草沼泽化草甸深层（10—20 cm）根系碳氮比。这些结果预示着增温使得土壤水分较低的小嵩草草甸朝着资源保守的慢速生长型发展,以适应暖干化的环境;土壤水分较高的藏嵩草沼泽化草甸朝着资源获取的快速生长型发展,加速利用土壤中的养分满足植物生长需要。可见,土壤水分可以调节高寒草甸对气候变暖的演变趋势,强调了水分的重要性。     

Simulated climate change: a field manipulation study of polar microarthropod community response to global warming

Passive cloches were deployed at three altitudinally distinct sites on Signy Island, maritime Antarctica, to investigate the effect of ameliorated thermal environment upon fellfield microarthropod communities Temperature was monitored at 1 5 m height, at ground surface level, and at 5 cm depth in cloche and control plots During summer (December - March), cloches elevated monthly mean temperatures by up to 2 46°C at the soil surface and 2 20°C at 5 cm depth Integrated air temperatures over consecutive 10 d periods were up to 4 65°C wanner in cloches than controls During winter (April - November), snow cover of the fellfield sites buffered temperature variation and reduced the treatment effect After eight years of these manipulations, sampling of the upper 50 mm of soil revealed consistently greater microarthropod populations within cloches than in controls (treatment effect p<0.05) Maximum difference occurred at high altitude where thermal amelioration was greatest (site effect p<0.05) Cloche populations of the numerically dominant collembolan Cryptopygus antarcticus Willem contained an increased proportion of small (length < 750 μm) individuals No species new to Signy Island were recorded Relating these microarthropod populations to the ameliorated thermal environment suggests that Antarctic invertebrate communities may respond to global warming, as predicted by global circulation models, with an increase in abundance with little increase in diversity However, this response could be indirect, the intermediate controlling factor being the percentage cover of the soil surface by vegetation, itself a function of climate change     

Extreme Weather Event Triggers Cascade Towards Extreme Turbidity in a Clear-water Lake

Climate forecasts project a global increase in extreme weather events, but information on the consequences for ecosystems is scarce. Of particular significance for lakes are severe storms that can influence biogeochemical processes and biological communities by disrupting the vertical thermal structure during periods of stratification. An exceptional storm passing over northern Germany in July 2011 provided an opportunity to assess the consequences and underlying mechanisms of such extreme events on the interplay between the physics and ecological characteristics of a deep, nutrient-poor lake. Wind speeds were among the most extreme on record. A suite of variables measured throughout the event consistently indicates that a cascade of processes pushed the clear-water lake into an exceptionally turbid state. Specifically, thermocline deepening by the storm-entrained cyanobacteria of a deep chlorophyll maximum located at about 8 m depth into the surface mixed layer. Released from light limitation, intense photosynthesis of the cyanobacteria boosted primary production, increased algal biomass, raised the pH and thus induced massive calcite precipitation to a level never observed within three decades of lake monitoring. As a consequence, water transparency dropped from 6.5 to 2.1 m, the minimum on record for 40 years, and the euphotic zone shrank by about 8 m for several weeks. These results show that cyanobacterial blooms not only are promoted by climate warming, but can also be triggered by extreme storms. Clear-water lakes developing a deep chlorophyll maximum appear to be particularly at risk in the future, if such events become more intense or frequent.