Summer use of a small stream by fish and crayfish and exchanges with adjacent lentic macrohabitats

1. Movements between a stream reach and two adjacent lentic macrohabitats, a beaver pond and a lake, were followed for the Appalachian crayfish and two fish species, brook charr and brown bullhead, over an 85‐d period from early June to late August, and were analysed in relation to water level, maximum water temperature, photoperiod length, lunar luminosity, and age, by use of time‐series regressions. 2. Brook charr showed strong net immigration to the stream reach for underyearling (age class 0+) fish but net emigration for 1+ fish. Both immigration and emigration were positively related to water level and temperature; migratory responses to temperature were age‐specific. 3. Brown bullhead used the stream primarily as a corridor for downstream migration from the beaver pond to the lake. As with brook charr, water level and temperature had a positive effect on movement and responses were stronger in younger individuals. 4. Crayfish emigrated from the stream during the summer. Movements were positively related to increases in water level and temperature, with responses of 1+ crayfish much stronger than those of older individuals. 5. The results indicate that the stream tributary played different roles for brook charr (nursery), brown bullhead (dispersal corridor) and Appalachian crayfish (early summer refuge), and suggest that variation in water level or temperature resulting from climatic change or local anthropogenic activities might modify connectivity between macrohabitats, particularly for younger individuals.     

Growth and food of young-of-the-year brook charr,Salvelinus fontinalis,in lake and creek environments

Synopsis Growth and diet of brook charr,Salvelinus fontinalis, during their first weeks of exogenous feeding were compared between the nearshore zone of a central Ontario lake and its small, inlet creeks. Food selection was related to size and age of charr and possibly availability of food items. Mean growth was similar between habitats despite differences in consumed food types and caloric values. Differences in social behaviour may have been responsible for different patterns of growth between habitats. The importance of creek habitats to the dynamics and fitness of lake populations of brook charr are suggested.     

Density,length distribution,and diet of age-0 arctic charrSalvelinus alpinus in the surf zone of Thingvallavatn,Iceland

Synopsis Population densities of age-0 arctic chaff in the surf zone averaged 1.83 and 4.70 fish m^-2 in August 1984 and June 1985, respectively. Length variation of the littoral fish was low in early summer, increasing in late summer and autumn. Newly emerged charr, ∼ 20 mm long, appeared in the stony shallow water zone during both May and June. From length variation and variation in mouth position of the young charr, it is concluded that at least two of the four chaff morphs in the lake are present in the surf zone during spring and summer. In August, some of the larger age-0 charr had moved out from the surf zone, into the pelagic and the deeper epibenthic waters. The food of young littoral charr was dominated by large chironomid larvae (instar 3 and 4) and pupae Contribution from the Thingvallavatn project     

The influence of groundwater withdrawal and land use changes on brook charr (<Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>) thermal habitat in two coldwater tributaries in Michigan,U.S.A.

Brook charr (Salvelinus fontinalis) is a sentinel fish species which requires clean, cold water habitats. As such, many jurisdictions in the United States where brook charr are present use this species as an indicator of ecosystem health. In Michigan, groundwater-dominated streams are currently being impacted by increased groundwater withdrawal and land use/land cover changes which alter stream temperatures and their flow, and thus have the potential to significantly influence brook charr production and behavior. We quantified the influence of groundwater withdrawal and land use alteration on thermal habitat availability for brook charr using a groundwater modeling tool that estimated changes in baseflow to a stream segment based on changes to the groundwater system due to groundwater withdrawal and changes in rates of recharge related to landscape changes within a watershed. Projected stream temperature changes were calculated using a stream temperature modeling tool and compared to the range of temperature preferenda for brook charr in order to evaluate the potential impact of policy decisions regarding water extraction and land use/cover changes. The models predicted relatively small changes in both stream baseflow and consequently, stream temperature, with increased groundwater withdrawal rates. Land use/land cover alterations which we analyzed were shown to either mitigate or enhance the loss of brook charr thermal habitat as a result of groundwater withdrawal, depending on its relationship to recharge dynamics. This study emphasizes the importance of collaboration between water, land, and fisheries managers to ensure brook charr population viability, productivity, and sustainability in the face of environmental change, increasing water use and development in the watershed.     

Habitat, diet and food assimilation of Arctic charr under the winter ice in two subarctic lakes

The Arctic charr Salvelinus alpinus populations of the subarctic lakes Takvatn and Fjellfrøsvatn, north Norway, concentrated in the littoral zones (0–15 m) of the lakes during the entire winter (December to May) despite very low temperatures (0·2 and 0·7° C). High prey availability, low predation and competition and comparatively better light under snow and ice in shallow compared with deep water are probable reasons. At ice break in June, all Arctic charr moved to the profundal zone for a brief period, probably in response to the sudden light increase and a profundal resource peak of chironomid pupae. In the summer, the Arctic charr are found in the pelagic, profundal and littoral zones of the lakes. These populations therefore perform regular habitat shifts between the littoral zone in the winter, the profundal zone at ice break and the whole lake in the summer and autumn. The fish fed continuously during winter despite the cold water and the poor light. Amphipods and chironomid larvae dominated the diet. Catch per unit effort, numbers of stomachs with food and food intake rates varied with the subarctic light cycle but were lowest after the winter solstice. The winter assimilation of energy was about equal to the standard metabolism in Takvatn but was higher in Fjellfrøsvatn. The assimilation increased in both lakes under the spring ice in May. The habitat choice, diet and energy assimilation indicate that the Arctic charr is well adapted to the extreme winter conditions of subarctic lakes.     

Emergence chronology of brook charr,Salvenus fontinalis,alevins in an acidic stream

Synopsis The emergence chronology of brook charr alevins in the Chikanishing River, Ontario, Canada and the concurrent changes in stream chemistry resulting from acidic snowmelt runoff were examined during the spring, 1986–1988. Emergence patterns were similar among years with the most intense emergence (> 70%) occurring during the declining stream discharge following the spring flood. This period coincided with stream pH depressions (minimum pH 5.2) in 2 of 3 years, suggesting emergence behaviour frequently subjects the earliest free-swimming life interval of brook charr to episodic acidification events.     

Growth and mortality of Arctic charr and European whitefish reared at low temperatures

This comparative study explores how low temperatures affect the mortality and growth of first generation hatchery-reared progeny of subarctic populations of Arctic charr (Salvelinus alpinus L.) and European whitefish (Coregonus lavaretus L.). Replicate fish groups where held under simulated natural light regimes (70°N) at three constant temperatures (1, 3 and 6°C). The mortality of Arctic charr was low (≤1.4%) at all temperature treatments, whereas the mortality of whitefish increased with decreasing temperature from 6% at 6°C to 33% at 1°C. The Arctic charr exhibited higher growth rates than whitefish at all three temperature regimes. All groups of Arctic charr increased in weight, whereas whitefish held at 1°C did not gain weight throughout the experimental period of 133 days. Arctic charr exhibited a large intraspecific variability in growth leading to large variations in size-structure, whereas whitefish in contrast showed very homogenous growth and size-structure patterns; a dissimilarity probably related to species-specific differences in antagonistic behaviour. Evidently, Arctic charr are more cold water adapted than whitefish and are able to maintain growth at extremely low temperatures. Arctic charr thus appear to be the most suitable species for aquaculture at low water temperatures.     

Seasonal variation in condition, growth and food habits of walleye in a Great Plains reservoir and simulated effects of an altered thermal regime

Catch rates in gillnets and relative weight ( W _r) of walleye Stizostedion vitreum , in Glen Elder Reservoir, Kansas, were lowest during the summer (June–August) and highest during the autumn (September–November). Approximately 80% of their annual growth in length and mass was attained during late summer and autumn. Growth was minimal during winter (January–February) and spring (March–May). The number of walleye with empty stomachs was highest during the summer. Invertebrates (Cladocera, Chironomidae) were common in walleye stomachs during the summer and spring, but contributed little to the ingested biomass. Gizzard shad Dorosoma cepedianum dominated walleye diets (per cent by mass) throughout the year. A bioenergetics model predicted that the proportion of maximum consumption ( P _c) was highest during the autumn and was probably due to spatial overlap of walleye and gizzard shad once water temperatures were <22° C. The bioenergetics model predicted that walleye would lose up to 65% of their body mass during the summer if water temperature increased by 10% (as predicted by some global warming models). Growth during the autumn, winter and spring was enhanced up to 150% by increased temperatures. The results of this study indicate that lower condition, reduced consumption and slow growth are a generalized response of walleye to extreme temperatures. Elevated temperatures may have a net positive effect on walleye growth if they can survive the high thermal stress during summer.     

Development of a species-specific fractionation equation for Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis> (L.)): an experimental approach

A species-specific fractionation equation for Arctic charr (Salvelinus alpinus (L.)) was developed experimentally for use in ecological studies of temperature-driven phenologies for the species. Juvenile Arctic charr were reared in controlled conditions at different temperatures (2–14°C), with three replicates of each temperature. Otoliths from the fish and water samples from the chambers were analysed for oxygen isotope composition and used to estimate temperature-dependent fractionation equations relating the isotopic ratio to rearing temperature. A linear and a second order polynomial relationship were estimated and validated using comparable Arctic charr data from another study. Temperatures predicted using the polynomial equation were not significantly different from recorded experimental temperatures, whereas with the linear equation there were significant differences between the predicted and measured temperatures. The polynomial equation also showed the least bias as measured by mean predictive error. Statistical comparisons of the polynomial fractionation equation to a similarly estimated equation for brook charr (Salvelinus fontinalis (Mitchill)) indicated significant differences. Results imply the need for species-specific fractionation equations, even for closely related fish. Results further suggest the polynomial form of the fractionation equation will facilitate more accurate characterisation of water temperatures suitable for use in ecological studies of temperature-driven phenologies of Arctic charr.     

Relationships between lake ecology and morphological characters in Icelandic Arctic charr,Salvelinus alpinus

The common occurrence of parallel phenotypic patterns suggests that a strong relationship exists between ecological dynamics and micro‐evolution. Comparative studies from a large number of populations under varying sets of ecological drivers could contribute to a better understanding of this relationship. We used data on morphology of arctic charr (Salvelinus alpinus) and ecological factors from 35 Icelandic lakes to test the hypothesis that morphological patterns among monomorphic charr populations from different lakes are related to interlake variation in ecological characteristics. There is extensive phenotypic diversity among populations of Icelandic charr, and populations are easily distinguished based on overall body morphology. The results obtained in the present study showed that the morphological diversity of charr was related to large‐scale diversity in lake ecology. Variation in charr morphology was related to water origin (e.g. spring fed versus run‐off), bedrock age, and fish community structure. The present study shows how various ecological factors can shape the biological diversity that we observe. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 761–771.     

Temperature impact on the midsummer decline of Daphnia galeata: an analysis of long-term data from the biomanipulated Bautzen Reservoir (Germany)

1. The influence of water temperature on occurrence and duration of a midsummer decline (MSD) of Daphnia galeata was studied in the biomanipulated Bautzen Reservoir in Germany. The proportion of piscivores in the fish community of the reservoir has been enhanced experimentally since 1981. As a consequence, Daphnia galeata has dominated the zooplankton. Over 18 years of study (1981–1998), a long‐lasting MSD (longer than 30 days) occurred in 7 years, whereas a short MSD (shorter than 30 days) was observed in 6 years. During the remaining 5 years, an MSD was not observed.
2. Two hypotheses were examined to explain the observed patterns. First, we postulated that high water temperature during winter and early spring (January–April) leads to an MSD after an early and high spring peak of daphnids. On the other hand, low temperature during winter and early spring should not cause an MSD owing to a slower increase of the population, resulting in a later peak of daphnids. Second, we hypothesized that the mean water temperature during early summer (May and June) influences the occurrence of an MSD (by controlling young‐of‐the‐year (YOY) fish predation on daphnids).
3. The water temperature during winter and early spring explains 83%, and the early summer water temperature 55%, of interannual variation in the occurrence of an MSD.
4. The interannual variation in duration of an MSD was neither explained by temperature during winter and early spring nor by early summer temperature alone, but in 14 of the 18 years (78%) by a combination of both.
5. We conclude that water temperature during winter and early spring had a strong impact on Daphnia mortality by influencing height and timing of the spring peak which, in turn, influenced the extent of overexploitation of their food resources. By contrast, the water temperature during early summer probably influenced the mortality of daphnids caused by predation of YOY fish. The relative timing of both sources of mortality, which depends on the temperature regime during the first 6 months of the year, is the key process in controlling the occurrence and duration of an MSD. A long‐lasting MSD, therefore, is likely in Bautzen Reservoir only if temperatures are high during winter and early spring, as well as during early summer.
6. As a consequence of climate warming, recent climate records reveal warming during winter, spring and early summer in middle Europe, rather than an increase in mean annual temperatures. If our findings and conclusions are related to this regional and temporal pattern of climate warming, an increasing frequency of years with a long‐lasting MSD and, consequently, a decreasing efficiency of biomanipulation can be predicted.     

Age, growth and reproduction of the cyprinid Rutilus lemmingii (Steindachner, 1866) in the River Huebra, Central Spain

Age, growth and reproduction of the R. lemmingii population of the River Huebra, Duero basin are analysed. Females dominated older age classes and lived up to 6 years (5+) while males only reached 5 years (4+). The growing season extended from April to September; growth rates were similar for both sexes. 0+ fish condition increased during their first summer and spring. In older fish, condition cycle was related to gonad development and showed some differences between sexes. The number of females reaching maturity at age 1 + almost doubled that of males; females also matured at smaller size. Both fecundity and egg size increased with female length: mean egg counts varied between 974 for 1 + individuals and an estimated 10491 for 5+ fish. Eggs were produced as a single batch, but were released fractionally during April and May.     

Context‐specific influence of water temperature on brook trout growth rates in the field

1. Modelling the effects of climate change on freshwater fishes requires robust field‐based estimates accounting for interactions among multiple factors. 2. We used data from an 8‐year individual‐based study of a wild brook trout (Salvelinus fontinalis) population to test the influence of water temperature on season‐specific growth in the context of variation in other environmental (i.e. season, stream flow) or biotic factors (local brook trout biomass density and fish age and size) in West Brook, a third‐order stream in western Massachusetts, U.S.A. 3. Changes in ambient temperature influenced individual growth rates. In general, higher temperatures were associated with higher growth rates in winter and spring and lower growth rates in summer and autumn. However, the effect of temperature on growth was strongly context‐dependent, differing in both magnitude and direction as a function of season, stream flow and fish biomass density. 4. We found that stream flow and temperature had strong and complex interactive effects on trout growth. At the coldest temperatures (in winter), high stream flows were associated with reduced trout growth rates. During spring and autumn and in typical summers (when water temperatures were close to growth optima), higher flows were associated with increased growth rates. In addition, the effect of flow at a given temperature (the flow‐temperature interaction) differed among seasons. 5. Trout density negatively affected growth rate and had strong interactions with temperature in two of four seasons (i.e. spring and summer) with greater negative effects at high temperatures. 6. Our study provided robust, integrative field‐based estimates of the effects of temperature on growth rates for a species which serves as a model organism for cold‐water adapted ectotherms facing the consequences of environmental change. Results of the study strongly suggest that failure to derive season‐specific estimates, or to explicitly consider interactions with flow regime and fish density, will seriously compromise our ability to predict the effects of climate change on stream fish growth rates. Further, the concordance we found between empirical observations and likely energetic mechanisms suggests that our general results should be relevant at broader spatial and temporal scales.     

Species-specific changes in the phenology and peak abundance of freshwater copepods in response to warm summers

SUMMARY 1. Climate warming is now widely recognised as a major factor influencing ecological processes in terrestrial, marine and freshwater habitats. Here, we investigated how a recent period of warm springs and summers has affected the population dynamics of various cyclopoid copepods in a central European lake. We compared (i) the duration of the period when the species were present in the water column, and (ii) their annual peak density in a period dominated by cool summers (1980–91) and one dominated by warm summers (1992–99). 2. The copepods under investigation were (i) Thermocyclops oithonoides, (ii) Mesocyclops leuckarti and (iii) Acanthocyclops robustus. These species differ in their thermal demand and seasonal phenology. Therefore, we hypothesised that enhanced summer warming would produce species‐specific responses. 3. The active phase of the copepods was usually prolonged both in spring and autumn. The earlier emergence of T. oithonoides (May in the warm years, July in the cool years) was probably related to high water temperature in late spring. The later onset of winter diapause in all species may have been coupled to raised temperature in late summer and autumn. 4. The annual peak abundance of the two thermophiles M. leuckarti and T. oithonoides increased significantly in the warm period. In the latter case, the increase was probably because of the early start to population growth. In contrast, M. leuckarti probably responded primarily to mid‐summer heat waves, in that its development time was likely to be short. We speculate that the increase in population size of both species resulted from the development of an additional generation (three instead of two cohorts per year). In contrast to these thermophiles, the coexisting A. robustus, which is adapted to a broader temperature range, did not respond noticeably to the warming trend. 5. In general, the nature of these responses to summer warming varied substantially among species, and depended on the detailed seasonal patterning of the warming. Our findings thus support the hypotheses that single species are sensitive indicators of climate change, and that the seasonal timing of warming is crucial in the context of climate–ecosystem relationships. 6. Moreover, our results add to the body of evidence that climate warming produces shifts in the seasonal phenology of aquatic and terrestrial organisms.     

Spring climate and summer otolith growth in juvenile Arctic charr, Salvelinus alpinus

The influence of different climate variables on the first four years of otolith growth in Salangen Arctic charr, Salvelinus alpinus, was studied over the period 1939?C2005. Salangen is a coastal, low altitude, subarctic lake system located in northern Norway. Climate data, including water temperature, air temperature, ice-cover and precipitation, were available for most of the 67?year period. Water temperatures in May and June had a significant effect on otolith growth during the second growth year, while no relationship between otolith growth and climate variables was found for the first, third and fourth years of otolith growth. Otolith increment size during the third and fourth growth year was autocorrelated with growth during the previous year. Spring snow fall and timing of ice break-up had an indirect effect on growth, as these variables were highly correlated with spring water temperatures. High variation in otolith growth within years and among individuals suggests that individual and age-specific variations in spatial habitat use may confound the direct effects of changing air temperatures and time of ice break-up.     

Winter ecology of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in a subarctic lake,Norway

We studied habitat choice, diet, food consumption and somatic growth of Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) during the ice-covered winter period of a subarctic lake in northern Norway. Both Arctic charr and brown trout predominantly used the littoral zone during winter time. Despite very cold winter conditions (water temperature <1°C) and poor light conditions, both fish species fed continuously during the ice-covered period, although at a much lower rate than during the summer season. No somatic growth could be detected during the ice-covered winter period and the condition factor of both species significantly declined, suggesting that the winter feeding rates were similar to or below the maintenance requirements. Also, the species richness and diversity of ingested prey largely decreased from summer to winter for both fish species. The winter diet of Arctic charr <20 cm was dominated by benthic insect larvae, chironomids in particular, and Gammarus lacustris, but zooplankton was also important in December. G. lacustris was the dominant prey of charr >20 cm. The winter diet of brown trout <20 cm was dominated by insect larvae, whereas large-sized trout mainly was piscivorous, feeding on juvenile Arctic charr. Piscivorous feeding behaviour of trout was in contrast rarely seen during the summer months when their encounter with potential fish prey was rare as the small-sized charr mainly inhabited the profundal. The study demonstrated large differences in the ecology and interactions of Arctic charr and brown trout between the winter and summer seasons.     

Production of brook charr and ouananiche in a large rapid,tributary of the Caniapiscau River,northern Quebec

Synopsis The age structures of brook charr (Salvelinus fontinalis) and ouananiche (Salmo salar) stocks inhabiting a large rapid the river Méo, tributary to the Caniapiscau River were used to compare population stability and production of these species in north central Quebec. The brook chart stock was stable whereas ouananiche showed considerable variation in year class strength. Stock estimates were not significantly different for the two species although the brook charr estimate was 1.5 that of the ouananiche. Production estimates differed by a greater margin because of different growth patterns. Brook chair production was estimated at 19.4 kg ha⁻¹ yr⁻¹. Above age 2+ it was 11.4 kg ha⁻¹ yr⁻¹ which compares with 4.8 kg ha⁻¹ yr⁻¹ for the same age groups of ouananiche.     

Water temperature, light intensity and zooplankton density and the feeding activity of juvenile brook charr (Salvelinus fontinalis)

SUMMARY 1. The objective of this study was to evaluate the effects of zooplankton biomass (as a measure of density), fish biomass, light intensity and water temperature on the attack rate and swimming characteristics (i.e. swimming speed and angle of turn) of juvenile (1+) brook charr (Salvelinus fontinalis) in field enclosures. We used a portable underwater camera system in a series of pelagic enclosures to quantify the feeding behaviour of brook charr over a gradient of natural conditions. 2. In simple linear or non‐linear regression models we found (i) that attack rate and angle of turn were positively related to water temperature, (ii) that attack rate and swimming speed were positively related to zooplankton biomass and light intensity and (iii) that attack rate was positively related to swimming speed. In multiple regression models, fish biomass, light intensity and variance of the angle of turn accounted for 87% of the variation in attack rate. Light intensity and water temperature accounted for 86% of the variation in swimming speed. Fish gut fullness and attack rate accounted for 83% of the variation in the variance of the angle of turn executed by fish. 3. The increase in the number of attacks as zooplankton biomass increases conforms to the general positive functional response observed in other fish species. Our results also support the hypothesis that swimming speed increases with prey biomass. We did not observe a plateau in attack rate as zooplankton biomass increased. As our experiments were performed under natural biotic and abiotic conditions, factors other than zooplankton biomass might affect or limit this response, such as water temperature and light intensity. 4. Because zooplankton biomass was correlated with water temperature and light intensity, it was not possible to evaluate the independent contribution of these factors on the attack rate and swimming characteristics (swimming speeds and angle of turn) of brook charr. However, this study highlighted the impact of these factors on the feeding behaviour of juvenile brook charr when feeding in the pelagic habitat under natural conditions, and their importance in future models of optimal foraging and fish habitat quality.     

An atypical brook charr (Salvelinus fontinalis) spawning area

Synopsis Brook charr (Salvelinus fontinalis) spawned successfully on a 0.3 m thick aggregation of waterlogged sticks, woodchips and debris overlying the soft ooze bottom of a small Precambrian Shield lake. Brook charr were apparently attracted to groundwater seeping up through the aggregation and utilized the tangle of various sized sticks as a spawning substrate. Eggs were deposited in late October and alevins emerged from the area in late March, periods that coincide with the conventional spawning of native brook charr on gravel areas in nearby lakes. Our observations support the contention that upwellin, water is more important than bottom type in stimulating brook charr to select a spawning site.     

The recovery of a very shallow eutrophic lake, 20 years after the control of effluent derived phosphorus

1. Monitoring at fortnightly to monthly intervals of a very shallow, lowland lake over 24 years has enabled the time course of recovery from nutrient enrichment to be investigated after high external P loading of the lake (>10 g P m^?2 year^?1) was reduced between 1977 and 1980. 2. The lake showed a relatively rapid response during the spring and early summer, with a reduction in phytoplankton biomass occurring after 5 years when soluble reactive phosphorus concentration was <10 μg L^?1. 3. However, during the later summer the response was delayed for 15 years because of sustained remobilisation of phosphorus from the sediment. The greater water clarity in spring and a gradual shift from planktonic to benthic algal growth may be related to the reduction in internal loading after 15 years. 4. Changes in the phytoplankton community composition were also observed. Centric diatoms became less dominant in the spring, and the summer cyanobacteria populations originally dominated by non‐heterocystous species (Limnothrix/Planktothrix spp.) almost disappeared. Heterocystous species (Anabaena spp. and Aphanizomenon flos‐aquae) were slower to decline, but after 20 years the phytoplankton community was no longer dominated by cyanobacteria. 5. There were no substantial changes in food web structure following re‐oligotrophication. Total zooplankton biomass decreased but body size of Daphnia hyalina, the largest zooplankton species in the lake, remained unchanged, suggesting that the fish population remained dominated by planktivorous species. 6. Macrophyte growth was still largely absent after 20 years, although during the spring water clarity may have become sufficient for macrophytes to re‐establish.