Seasonal influence of brook trout and mottled sculpin on lower trophic levels in an Appalachian stream

1. In some situations fish have strong top‐down effects in stream communities while in others they seem to be relatively unimportant. Differences in the impact of fish may depend on a variety of factors including the foraging mode of the fish, interactions among fish species and temporal variation in environmental conditions and species interactions. 2. We investigated the effect of brook trout (Salvelinus fontinalis) and mottled sculpin (Cottus bairdi) on lower trophic levels in Appalachian streams and whether or not interactions between these fish changed their influence. Mesocosms were placed in a headwater stream in a randomized complete block design. Within blocks, mesocosms were randomly assigned to one of the following treatments: (i) no fish; (ii) sculpin only; (iii) trout only and (iv) both sculpin and trout. Fish biomass was the same in all three fish treatments. Invertebrate density and algal biomass in mesocosms were determined after 3 weeks. We repeated the experiment in the autumn, spring and summer to test for seasonality of fish effects. 3. The effect of fish on invertebrate assemblages was seasonal and depended on prey identity. Sculpin strongly suppressed grazer abundance in spring while trout had little effect on grazers in any season. The influence of both fish on insect predators was similar and relatively constant across seasons. We found little evidence of an interaction between sculpin and trout that strongly influenced their effect on prey across seasons. 4. None of the fish treatments influenced algal biomass during any of the seasons. Algal growth was also seasonal, with a two‐ to four‐fold increase in algal biomass in spring compared to autumn and summer. 5. Our results indicate that benthic and drift feeding fish differ in their effects on some, but not all prey. Furthermore, fish effects on prey were strongly seasonal for some, but not all prey types. While the temporal context is not commonly considered, our results indicate seasonality can be an important component of predator–prey interactions in streams.     

Changes in seasonal climate outpace compensatory density‐dependence in eastern brook trout

Understanding how multiple extrinsic (density‐independent) factors and intrinsic (density‐dependent) mechanisms influence population dynamics has become increasingly urgent in the face of rapidly changing climates. It is particularly unclear how multiple extrinsic factors with contrasting effects among seasons are related to declines in population numbers and changes in mean body size and whether there is a strong role for density‐dependence. The primary goal of this study was to identify the roles of seasonal variation in climate driven environmental direct effects (mean stream flow and temperature) vs. density‐dependence on population size and mean body size in eastern brook trout (Salvelinus fontinalis). We use data from a 10‐year capture‐mark‐recapture study of eastern brook trout in four streams in Western Massachusetts, USA to parameterize a discrete‐time population projection model. The model integrates matrix modeling techniques used to characterize discrete population structures (age, habitat type, and season) with integral projection models (IPMs) that characterize demographic rates as continuous functions of organismal traits (in this case body size). Using both stochastic and deterministic analyses we show that decreases in population size are due to changes in stream flow and temperature and that these changes are larger than what can be compensated for through density‐dependent responses. We also show that the declines are due mostly to increasing mean stream temperatures decreasing the survival of the youngest age class. In contrast, increases in mean body size over the same period are the result of indirect changes in density with a lesser direct role of climate‐driven environmental change.     

Interacting effects of temperature and density on individual growth performance in a wild population of brown trout

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Trout affect the density, activity and feeding of a larval plethodontid salamander

1. Ecologists have struggled to describe general patterns in the impacts of predators on stream prey, particularly at large, realistic spatial and temporal scales. Among the confounding variables in many systems is the presence of multiple predators whose interactions can be complex and unpredictable. 2. We studied the interactions between brook trout (Salvelinus fontinalis) and larval two‐lined salamanders (Eurycea bislineata), two dominant vertebrate predators in New England stream systems, by examining patterns of two‐lined salamander abundance in stream reaches above and below waterfalls that are barriers to fish dispersal, by measuring the effects of trout on salamander density and activity using a large‐scale manipulation of brook trout presence, and by conducting a small‐scale laboratory experiment to study how brook trout and larval two‐lined salamanders affect each other's prey consumption. 3. We captured more salamanders above waterfalls, in the absence of trout, than below waterfalls where trout were present. Salamander density and daytime activity decreased following trout addition to streams, and salamander activity shifted from aperiodic to more nocturnal with fish. Analysis of stomach contents from our laboratory experiment revealed that salamanders eat fewer prey with trout, but trout eat more prey in the presence of salamanders. 4. We suggest that as predators in streams, salamanders can influence invertebrate prey communities both directly and through density‐ and trait‐mediated interactions with other predators.     

Non‐additive effects of density‐dependent and density‐independent factors on brown trout vital rates

Interactions between density‐dependent and density‐independent processes can lead to variation in both growth and survival rates. Detecting such effects, however, will often require sampling on an individual level and at the appropriate spatial and temporal scale. This study documents substantial variation in survival and growth of stream‐dwelling brown trout Salmo trutta from a small Norwegian stream. The data is based on seasonal capture–recaptures of individually marked trout on fixed stations during eight years. The fish were small‐sized, rarely reaching sizes larger than 20 cm and ages older than seven years. Density varied between 0.2–0.8 fish m^?2. Variation in survival and recapture probabilities was analysed using program MARK. Apparent survival (the probability of being alive and present within the study area) generally decreased with increasing trout density and increasing drought level (measured as lowest observed water flow) during both winter and summer. Further, there was a significant interaction effect between density and water flow, indicating that density‐dependent effects on survival predominated when environmental conditions were benign (no drought), while density‐independent processes were most important under harsh environmental conditions (drought). Observed length‐at‐age during autumn indicated a more or less linear growth trajectory throughout life, and no effect of density, water flow or temperature was found. However, using the individual‐based capture–recapture data to estimated specific growth rate, significant positive effects of water flow and temperature and a negative effect of density were identified. Thus, the capture–recapture data suggest a strong potential for population regulation at the rather low densities found in this stream, and regulation may occur both through effects on survival and growth.     

A bioenergetic assessment of the influence of stocking practices on rainbow trout (Oncorhynchus mykiss) growth and consumption in a New Zealand lake

Summary

• 1 To investigate the carrying capacity and factors affecting growth of rainbow trout in Lake Rotoiti, we employed a bioenergetics model to assess the influence of stocking rates, timing of releases and prey abundance on growth and prey consumption. We hypothesised that stocking rates and prey abundance would affect growth and prey consumption by influencing per‐capita prey availability, and that the environmental conditions encountered by fish at the time of stocking would affect growth and consumption.
• 2 Prey consumption of stocked rainbow trout was calculated with the Wisconsin bioenergetics model. We calculated growth trajectories of released trout based on data from stocked trout that were released in spring and autumn from 1993 to 2009 and then re‐captured by anglers. Diet, prey energy density, body mass lost during spawning and lake temperature were measured locally.
• 3 Stocking timing had no effect on return rates to anglers or length or weight of caught fish. Although trout released in autumn were smaller than those released in spring, autumn‐released trout grew at a faster rate and had similar lengths and weights to spring cohorts after 2 years of growth in the lake. Modelled consumption parameters were negatively correlated with trout population size, suggesting that stocking rates (347–809 fish ha^?1 year^?1) caused density‐dependent effects on growth. Although common smelt (Retropinna retropinna) accounted for 85% of total prey consumption, no significant relationship was found between prey consumption by individual trout and adult smelt abundance, possibly because trout are targeting smaller smelt that our abundance estimate did not account for.
• 4 Releasing trout in autumn appears to be advantageous for growth, possibly because (i) temperature is more suitable for growth in autumn–winter than in spring–summer and (ii) prey for small trout is abundant in autumn. Mild winter conditions appear to enhance overwinter survival and growth of rainbow trout in warm‐temperate lakes compared to higher latitudes. This implies that moderately productive warm‐temperate lake ecosystems are highly suitable for trout growth in winter, but less so in summer, when lake stratification and high nutrient levels may create conditions suitable for algal blooms and hypolimnetic deoxygenation. High growth rates of trout in warm‐temperate lakes can therefore be supported by timing releases to coincide with favourable winter conditions.

     

Seasonal temperature and precipitation regulate brook trout young‐of‐the‐year abundance and population dynamics

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Size‐dependent survival of brook trout Salvelinus fontinalis in summer: effects of water temperature and stream flow

A 5 year individual‐based data set was used to estimate size‐specific survival rates in a wild brook trout Salvelinus fontinalis population in a stream network encompassing a mainstem and three tributaries (1·5–6 m wetted width), western Massachusetts, U.S.A. The relationships between survival in summer and temperature and flow metrics derived from continuous monitoring data were then tested. Increased summer temperatures significantly reduced summer survival rates for S. fontinalis in almost all size classes in all four sites throughout the network. In contrast, extreme low summer flows reduced survival of large fish, but only in small tributaries, and had no significant effects on fish in smaller size classes in any location. These results provide direct evidence of a link between season‐specific survival and environmental factors likely to be affected by climate change and have important consequences for the management of both habitats and populations.     

Seasonal and regional infestation characteristics of three ectoparasites of whiting, Merlangim merlangus L., in the North Sea

The colonization by both resident and migrating spawner populations of brown trout and the characteristics of resident and migrating juveniles derived from the two populations have been studied in a brook and its tributary over 4 years. Resident trout spawns mainly in the upstream part of the brook and migrating trout in the downstream part. There are density and growth variations for the two age classes (0+ and 1 +) of juveniles in autumn according to the year and the environment. In the brook, the population of 0 + fish increases from downstream to upstream while the density of other age classes decreases. The migrating juvenile population of the brook changes annually and consists mainly of 1 s (one summer) individuals coming from the upper part. These individuals migrate generally in autumn and winter while young trout produced in the middle and downstream parts of the brook migrate mainly in the spring. The emigration process of the 0 + population decreases markedly from upstream to downstream and appears to be independent of the autumn length and sex ratio. In the tributary, most trout are 0+ years old, the population structure is different, and no migrating fish is observed. The results are discussed and a colonization strategy of the brown trout population in this brook is suggested.     

Growth rate differences between resident native brook trout and non-native brown trout

Between species and across season variation in growth was examined by tagging and recapturing individual brook trout Salvelinus fontinalis and brown trout Salmo trutta across seasons in a small stream (West Brook, Massachusetts, U.S.A.). Detailed information on body size and growth are presented to (1) test whether the two species differed in growth within seasons and (2) characterize the seasonal growth patterns for two age classes of each species. Growth differed between species in nearly half of the season- and age-specific comparisons. When growth differed, non-native brown trout grew faster than native brook trout in all but one comparison. Moreover, species differences were most pronounced when overall growth was high during the spring and early summer. These growth differences resulted in size asymmetries that were sustained over the duration of the study. A literature survey also indicated that non-native salmonids typically grow faster than native salmonids when the two occur in sympatry. Taken together, these results suggest that differences in growth are not uncommon for coexisting native and non-native salmonids.     

An investigation of the advantages of autumn and spring stocking with brown trout Salmo trutta L. in a Yorkshire reservoir

Batches of trout have been introduced into Chelker Reservoir in Yorkshire in the autumn and spring since the 1870's for angling purposes. Six batches of tagged, hatchery-reared brown trout Salmo trutta L. were introduced from autumn 1966 to spring 1969. During the angling season fish introduced in the spring give better catches than those stocked in the autumn. At the beginning of the season the larger fish in the spring batch are caught more often than the smaller fish from the same batch. The larger fish in the autumn batch are caught more often than the smaller fish from that batch throughout the season. The population, available to the angler from the shore was estimated to be 1491 in 1968, with 722 fish/km of shoreline. More fish survive to a second year in the reservoir than is apparent from the number of tags returned. Fish introduced in the spring usually begin growing before those introduced in the autumn, thereafter growth rates varied. The growth rate was independent of the number offish stocked up to the numbers put in.
Batches of tagged trout were retained at the hatchery up to nine months to gain relevant experience of post-tagging mortalities, tag loss rate and effect of tags on growth.     

Brown trout and food web interactions in a Minnesota stream

1. We examined indirect, community‐level interactions in a stream that contained non‐native brown trout (Salmo trutta Linnaeus), native brook trout (Salvelinus fontinalis Mitchill) and native slimy sculpin (Cottus cognatus Richardson). Our objectives were to examine benthic invertebrate composition and prey selection of fishes (measured by total invertebrate dry mass, dry mass of individual invertebrate taxa and relative proportion of invertebrate taxa in the benthos and diet) among treatments (no fish, juvenile brook trout alone, juvenile brown trout alone, sculpin with brook trout and sculpin with brown trout). 2. We assigned treatments to 1 m² enclosures/exclosures placed in riffles in Valley Creek, Minnesota, and conducted six experimental trials. We used three designs of fish densities (addition of trout to a constant number of sculpin with unequal numbers of trout and sculpin; addition of trout to a constant number of sculpin with equal numbers of trout and sculpin; and replacement of half the sculpin with an equal number of trout) to investigate the relative strength of interspecific versus intraspecific interactions. 3. Presence of fish (all three species, alone or in combined‐species treatments) was not associated with changes in total dry mass of benthic invertebrates or shifts in relative abundance of benthic invertebrate taxa, regardless of fish density design. 4. Brook trout and sculpin diets did not change when each species was alone compared with treatments of both species together. Likewise, we did not find evidence for shifts in brown trout or sculpin diets when each species was alone or together. 5. We suggest that native brook trout and non‐native brown trout fill similar niches in Valley Creek. We did not find evidence that either species had an effect on stream communities, potentially due to high invertebrate productivity in Valley Creek.     

Forests and the temperature of upland streams in Wales: a modelling exploration of the biological effects

SUMMARY. 1. Daily temperature data from six streams in upland Wales were used to explore the thermal effects of afforestation on stream ecology. The data were linked to published biological models to simulate fish and invertebrate development.
2. Mean daily temperatures in forest streams were lower than those of moorland streams in spring and summer, and higher in winter. These spatial comparisons were supported by the results of experimental bank-side clearance at a forest site, where there was evidence that stream temperatures fell in winter and rose in spring following treatment.
3. Simulations indicated that brown trout (Salmo trutta) could weigh over 30% more by the end of their second growing season in a moorland compared with a forest stream. Several species of insects showed slower simulated egg development at forest sites. For two ephemeropteran species simulated nymphal growth was also retarded, suggesting significant alterations to the life cycle. Two plecopteran species were affected only slightly by the different temperature regimes.
4. Overall, the simulations suggested that afforestation, by reducing summer temperatures, could lead to marked reductions in rates of development of some invertebrates and fish.     

Migration of anadromous brown trout Salmo trutta in a Norwegian river

1. Upstream and downstream migrating anadromous brown trout Salmo trutta were monitored daily in fish traps in the River Imsa in south-western Norway for 24 years, from 1976 to 1999. One-third of the fish descended to sea during spring (February–June) and two-thirds during autumn (September–January).
2. In spring, high water temperature appeared to influence the downstream descent. Large brown trout (> 30 cm, chiefly two or more sea sojourns) descended earlier and appeared less dependent on high water temperature than smaller and younger fish. The spring water flow was generally low and of little importance for the descent.
3. In autumn, the daily number of descending brown trout correlated positively with flow and negatively with water temperature.
4. Brown trout ascended from the sea between April and December, but more than 70% ascended between August and October. The number of ascending trout increased significantly with both decreasing temperature and flow during the autumn. This response to flow appeared to be the result of the autumn discharge which is generally high and most fish ascended at an intermediate flow of 7.5–10 m³ s⁻¹ (which is low for the season).
5. In a river like the Imsa with low spring and high autumn flows, water temperature appears to be the main environmental factor influencing the timing and rate of spring descent, while both water temperature and flow seemed to influence the timing and rate of the autumn descent and ascent. These relationships make sea trout migrations susceptible to variation in climate and human impacts of the flow regime in rivers.     

Population regulation of brook trout (Salvelinus fontinalis) in Hunt Creek,Michigan: a 50‐year study

1. Fisheries models generally are based on the concept that strong density dependence exists in fish populations. Nonetheless, there are few examples of long‐term density dependence in fish populations. 2. Using an information theoretical approach (AIC) with regression analyses, we examined the explanatory power of density dependence, flow and water temperature on the per capita rate of change and growth (annual mean total length) for the whole population, adults, 1+ and young‐of‐the‐year (YOY) brook trout (Salvelinus fontinalis) in Hunt Creek, Michigan, USA, between 1951 and 2001. This time series represents one of the longest quantitative population data sets for fishes. 3. Our analysis included four data sets: (i) Pooled (1951–2001), (ii) Fished (1951–65), (iii) Unfished (1966–2001) and (iv) Temperature (1982–2001). 4. Principle component analyses of winter flow data identified a gradient between years with high mean daily winter flows, high daily maximum and minimum flows and frequent high flow events, and years with an opposite set of flow characteristics. Flows were lower during the Fished Period than during the Unfished Period. Winter temperature analyses elucidated a gradient between warm mean, warm minimum and maximum daily stream temperatures and a high number of minimum daily temperatures >6.1 °C, and years with the opposite characteristics. Summer temperature analyses contrasted years with warm summer stream temperatures vs years with cool summer stream temperatures. 5. Both YOY and adult densities varied several‐fold during the study. Regression analysis did not detect a significant linear or nonlinear stock–recruitment relationship. AIC analysis indicated that density dependence was present in 15 of 16 cases (four population segments × four data sets) for both per capita rate of increase (w_i values 0.46–1.00) and growth data (w_i values 0.28–0.99). The almost ubiquitous presence of density dependence in both population and growth data is concordant with results from other trout populations and other studies in Michigan.     

The temperature–productivity squeeze: constraints on brook trout growth along an Appalachian river continuum

We tested the hypothesis that brook trout growth rates are controlled by a complex interaction of food availability, water temperature, and competitor density. We quantified trout diet, growth, and consumption in small headwater tributaries characterized as cold with low food and high trout density, larger tributaries characterized as cold with moderate food and moderate trout density, and large main stems characterized as warm with high food and low trout density. Brook trout consumption was highest in the main stem where diets shifted from insects in headwaters to fishes and crayfish in larger streams. Despite high water temperatures, trout growth rates also were consistently highest in the main stem, likely due to competitively dominant trout monopolizing thermal refugia. Temporal changes in trout density had a direct negative effect on brook trout growth rates. Our results suggest that competition for food constrains brook trout growth in small streams, but access to thermal refugia in productive main stem habitats enables dominant trout to supplement growth at a watershed scale. Brook trout conservation in this region should seek to relieve the “temperature–productivity squeeze,” whereby brook trout productivity is constrained by access to habitats that provide both suitable water temperature and sufficient prey.     

Can replacement of native by non‐native trout alter stream‐riparian food webs?

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Importance of terrestrial subsidies for native brook trout in Appalachian intermittent streams

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Influence of spring and summer water temperature on brook charr, Salvelinus fontinalis, growth and age structure in the Ford River, Michigan

Synopsis The influence of late spring and summer water temperatures on brook charr, Salvelinus fontinalis, growth and age structure was evaluated from 1984 to 1991 in the Ford River, Michigan. Temperature was monitored and brook charr sampled for vital statistics from late May through September using fyke nets and weirs at four locations within a 25.8 km section of stream. Scale analysis was used to determine captured brook charr age, past length at age and relative annual growth rates. Late spring and summer water temperature patterns varied between years with the greatest variability occurring in May and June. Age and size structure also varied between years and was significantly related to temperature. Years with cooler late spring and summer temperature patterns were dominated by older (age 2 and 3), larger brook charr, while years with warmer spring and summer temperature patterns were dominated by younger (age 1), smaller brook charr. Spring and summer temperature did not appear to have a significant effect on the growth of age 0 or age 1 brook charr. However, temperature was negatively related to brook charr growth from age 2 on. As spring and summer water temperatures are critical to brook charr growth and survival, it is important that a streams thermal regime be considered when establishing management goals for this species.     

Thermal controls of Yellowstone cutthroat trout and invasive fishes under climate change

We combine large observed data sets and dynamically downscaled climate data to explore historic and future (2050–2069) stream temperature changes over the topographically diverse Greater Yellowstone Ecosystem (elevation range = 824–4017 m). We link future stream temperatures with fish growth models to investigate how changing thermal regimes could influence the future distribution and persistence of native Yellowstone cutthroat trout (YCT) and competing invasive species. We find that stream temperatures during the recent decade (2000–2009) surpass the anomalously warm period of the 1930s. Climate simulations indicate air temperatures will warm by 1 °C to >3 °C over the Greater Yellowstone by mid‐21st century, resulting in concomitant increases in 2050–2069 peak stream temperatures and protracted periods of warming from May to September (MJJAS). Projected changes in thermal regimes during the MJJAS growing season modify the trajectories of daily growth rates at all elevations with pronounced growth during early and late summer. For high‐elevation populations, we find considerable increases in fish body mass attributable both to warming of cold‐water temperatures and to extended growing seasons. During peak July to August warming, mid‐21st century temperatures will cause periods of increased thermal stress, rendering some low‐elevation streams less suitable for YCT. The majority (80%) of sites currently inhabited by YCT, however, display minimal loss (<10%) or positive changes in total body mass by midcentury; we attribute this response to the fact that many low‐elevation populations of YCT have already been extirpated by historical changes in land use and invasions of non‐native species. Our results further suggest that benefits to YCT populations due to warmer stream temperatures at currently cold sites could be offset by the interspecific effects of corresponding growth of sympatric, non‐native species, underscoring the importance of developing climate adaptation strategies that reduce limiting factors such as non‐native species and habitat degradation.