Interspecific competition between stream-dwelling brown trout and Alpine bullhead

Density and composition of benthic invertebrates and the diet of brown trout Salmo trutta and Alpine bullhead Cottus poecilopus were studied at two sites in one Norwegian stream. The sites were separated by an impassable waterfall, and brown trout density was five to 10 times higher at the upper, allopatric site than downstream where it lived in sympatry with the Alpine bullhead. Benthic invertebrate communities did not differ between sites; however, the size distribution of chironomids and trichopterans were skewed towards lighter individuals at the sympatric site. Diet composition suggested that sympatric brown trout foraged more on invertebrate drift and from the surface than allopatric brown trout. Alpine bullhead diet did not differ significantly from brown trout diet, except that the Alpine bullhead fed on heavier individual prey within a few taxa and did not consume chironomid pupae or surface insects. The collected data support the hypothesis that brown trout living in sympatry with Alpine bullhead feed at locations with higher predation risk, which is a probable explanation for their lower population density.     

Effects of species invasion on population dynamics,vital rates and life histories of the native species

Invasions occurring in natural environments provide the opportunity to study how vital rates change and life histories evolve in the presence of a competing species. In this work, we estimate differences in reproductive traits, individual growth trajectories, survival, life histories and population dynamics between a native species living in allopatry and in sympatry with an invasive species of the same taxonomic Family. We used as a model system marble trout Salmo marmoratus (native species) and rainbow trout Oncorhynchus mykiss (non-native) living in the Idrijca River (Slovenia). An impassable waterfall separates the stream into two sectors only a few 100 meters apart: a downstream sector in which marble trout live in sympatry with rainbow trout and an upstream sector in which marble trout live in allopatry. We used an overarching modelling approach that uses tag-recapture and genetic data (>2,500 unique marble and rainbow trout were sampled and genotyped) to reconstruct pedigrees, test for synchrony of population dynamics and model survival and growth, while accounting for individual heterogeneity. The population dynamics of the two marble trout populations and of rainbow trout were synchronous. We found higher prevalence of younger parents, higher mortality and lower population density in marble trout living in sympatry with rainbow trout than in marble trout living in allopatry. There were no differences in the average individual growth trajectories between the two marble trout populations. Faster life histories of marble trout living in sympatry with rainbow trout are consistent with predictions of life history theory.     

Microgeographical variation in brown trout reproductive traits: possible effects of biotic interactions

This study documents substantial variation in reproductive traits among populations of stream-dwelling brown trout ( Salmo trutta L.) at a very small geographic scale. Within two streams, we found a parallel pattern of variation, where females living above major waterfalls produced fewer and larger eggs than conspecifics from below the waterfalls. Four additional streams were represented with either a below-waterfall site ( n =2) or an above-waterfall site ( n =2). When these streams were included in the analyses, there was no consistent difference in reproductive traits between females from above- and below-waterfall sites. There was no significant difference in total reproductive investment among sites within streams, but considerable variation among streams. Female first-year growth rates was estimated from scales, and differed significantly among populations. Within streams, females from below waterfalls experienced higher first-year growth rates as compared to females from above the waterfalls. Within seven out of eight populations, egg size increased significantly with increasing female body length. Within three populations, we found evidence for a trade-off between offspring size and offspring number, as a negative association between fecundity and egg size independently of adult body size. Within three populations egg size decreased significantly with increasing maternal first-year growth, independently of adult body size. We suggest that the within-stream differences in offspring size/number strategies are influenced by population density and growth effects. Earlier, we have shown that population densities are consistently lower below the waterfalls in these streams. The Alpine bullhead ( Cottus poecilopus ) is found only below the waterfalls and could influence brown trout demography.     

Habitat degradation and trout stocking can reinforce the impact of flash floods on headwater specialist Alpine bullhead Cottus poecilopus – A case study from the Carpathian Mountains

In freshwater streams, flooding is a typical source of natural disturbance that plays a key role in the dynamics of animal populations and communities. However, habitat degradation and fish stocking might increase the severity of its impact. We tested the effects of a flash flood on the abundance of three size classes of headwater dwelling Alpine bullhead, Cottus poecilopus, in the streams of the Carpathian Mountains in the Czech Republic, that are stocked with hatchery‐reared brown trout, Salmo trutta. We showed that the overall abundance of Alpine bullhead was highest at the sites with the least degraded habitat (i.e., natural habitat) and we caught almost no Alpine bullhead at the sites with the most degraded habitat. The flash flood had a strong negative effect on the abundance of the largest individuals of Alpine bullhead. Abundance of small and medium size Alpine bullhead was negatively affected by the abundance of adult stocked brown trout before as well as after the flash flood. However, negative effect of adult brown trout abundance on abundance of large Alpine bullhead was not significant before the flash flood, and it became significant after the flash flood. This could indicate an accumulation of negative impacts of trout stocking and flash flood on this size class. Overall, our results suggest that stocking of hatchery trout and habitat degradation can reinforce the impact of flash floods on the population of Alpine bullhead in the streams of the Carpathian Mountains.     

Spatial and temporal population structure of sea trout at the Island of Gotland, Sweden, delineated from mitochondrial DNA

In a study of the genetic relationships among 879 anadromous brown trout Salmo trutta from 13 streams at the Island of Gotland, Sweden, using RFLP analysis of a mitochondrial DNA segment (NADH dehydrogenase-1 gene), six haplotypes were detected. Significant genetic divergence was observed among streams as well as between cohorts within streams. Approximately 8–9% of the total variation was due to differences between populations, and 4–5% was explained by differences between cohorts within populations. The female effective population size ( N _ef) was assessed from temporal haplotype frequency differences between consecutive cohorts; the estimated average N _ef over all populations was just below 30, suggesting that these populations were effectively quite small. With such small effective sizes the populations are expected to loose genetic variability quickly, but the observed levels do not appear particularly low. This indicates that female migration between streams occurs. The observed level of differentiation does not support the presumption that a particular pre-smolt migratory behaviour observed in Gotland streams, with large portions of fry leaving for the sea soon after hatching, results in a reduced homing ability. From a conservation management perspective the Gotland brown trout streams should be regarded as a population system where the vitality and survival of brown trout in one stream is dependent on the opportunity of contact and exchange of individuals from other streams.     

The populations of fish in the River Tees system on the Moor House National Nature Reserve, Westmorland

The age, growth, population densities and annual production of an isolated population of brown trout ( Salmo trutta L.) and bullhead ( Coitus gobio L.) were investigated over a 5 year period in Trout Beck in the northern Pennines. Additional data were obtained from two other sites; Great Dodgen Pot Sike and the River Tees.
Trout growth rates were low, particularly in Dodgen Pot Sike, but ages up to VIII were recorded. Bullhead growth rates were similar to those in the Tees at Cow Green (Crisp et al. , 1974), though the Moor House bullheads had a higher survival rate. Population densities of trout ranged from 0.10–0.22/m², with a fry contribution of 0.80%. The bullhead population in Trout Beck was sparse, generally 0.1/m², though an 0 group value of 0.46/m^a was estimated in 1967. At Tees Bridge values ranged from 2.5–7/8/m². Most trout matured by age III, though in Great Dodgen Pot Sike only half of the females were mature at age IV.
Annual production was estimated from Allen graphs. Trout production ranged from 1.02–3.50 g/m²/year, the fry contributing up to 51.5% of the total. A value of 0.48 g/m²/year was obtained for bullheads in the Trout Beck system in a year of good recruitment, whilst bullhead production at Tees Bridge was 7.43 g/m²/year, with fry and I group contributing 48.7 and 37.5% respectively. The high survival rate, irregular recruitment, and poor growth of these high altitude populations is discussed. Growth and reproductive peculiarities of the Great Dodgen Pot Sike trout are also discussed.     

Anserine and free amino acid contents in the egg and fry stages of developing rainbow trout (Oncorhynchus mykiss).

1. Anserine and free amino acid contents were examined in rainbow trout eggs, yolk sac and yolk-free fry before feeding. 2. Free amino acid levels showed little change in eggs and yolk sac and had a tendency to increase in yolk-free fry. 3. No anserine was detected in the eggs and yolk sac, but yolk-free fry significantly increased its concentration after hatching. This continuous increment of anserine was discussed in relation to buffering capacity of adult fish muscle.     

Recruitment variation in a stream galaxiid fish: multiple influences on fry dynamics in a heterogeneous environment

1. The relative importance of density‐dependent and density‐independent processes in explaining fluctuations in natural populations has been widely debated. In particular, the importance of larval supply and whether it may control the type of regulatory processes a population experiences has proved contentious. 2. Using surveys and field experiments conducted in streams in Canterbury, New Zealand, we investigated how variation in the survival of non‐migratory Galaxias vulgaris fry was affected by density‐dependent and density‐independent processes and how this variation influenced recruitment dynamics. 3. Fry populations with high settlement densities experienced a 70–80% reduction in population size from density‐related mortality during the first fourteen days after peak settlement but thereafter the influence of density‐dependent processes on fry was weak. The impact of environmental conditions on fry populations was dependent on fry size and the magnitude of the perturbation, such that flooding effects on fry survival were most severe when fry were small. 4. In streams not affected by flooding, the size and density of introduced trout (Salmo trutta and Oncorhynchus mykiss) were the most significant factors determining the abundance of eventual recruits. A field experiment manipulating brown trout access to fry populations revealed that trout as small as 110 mm may be capable of greatly reducing and possibly preventing galaxiid recruitment. 5. Overall, the results indicated density‐dependent population regulation was only possible at sites with high native fish densities because trout were likely to be suppressing the number of potential recruits at sites with low native fish numbers. Whilst density‐dependent processes had a strong effect on fry survival following the period of peak fry abundance, density‐independent processes associated with flow and predatory trout influences on fry survival largely determined recruitment variability among galaxiid populations. Focusing conservation efforts on improving habitat to increase fry retention and reducing the impacts of trout on galaxiids would ensure more native fish populations reached their potential abundance.     

Effects of sea trout stocking on the population genetics of landlocked brown trout, Salmo trutta L., in the Conwy River system, North Wales, U.K.

The effect of the introduction of fry of anadromous sea trout, Salmo trutta L., on the genetic integrity of landlocked brown trout populations was evaluated. Samples were taken from six brown trout populations from streams above impassable waterfalls in the Conwy river system (North Wales, U.K.) in 1989 and 1990. Three of these streams had no known stocking history and three had been stocked with sea trout fry from the lower Conwy system over the last few years. Representatives of these sea trout were collected from two streams in the lower Conwy system and from a hatchery. Allele frequencies at 13 loci, six of which were polymorphic, were determined by starch gel electrophoresis.
The stocked populations were intermediate in their allele frequencies between unstocked brown trout and sea trout samples. A principal component analysis suggested significant numbers of hybrids in all of the stocked streams. This shows that some of the introduced sea trout did not migrate down the falls to the sea, but stayed in fresh water and hybridized with the local population. The significance of this finding for the conservation of the genetic resource of brown trout stocks is discussed.     

Timing matters: species‐specific interactions between spawning time,substrate quality,and recruitment success in three salmonid species

Substratum quality and oxygen supply to the interstitial zone are crucial for the reproductive success of salmonid fishes. At present, degradation of spawning grounds due to fine sediment deposition and colmation are recognized as main factors for reproductive failure. In addition, changes in water temperatures due to climate change, damming, and cooling water inlets are predicted to reduce hatching success. We tested the hypothesis that the biological effects of habitat degradation depend strongly on the species‐specific spawning seasons and life‐history strategies (e.g., fall‐ vs. spring‐spawners, migratory vs. resident species) and assessed temperature as an important species‐specific factor for hatching success within river substratum. We studied the species‐specific differences in their responses to such disturbances using egg‐to‐fry survival of Danube Salmon (Hucho hucho), resident brown trout (Salmo trutta fario), and migratory brown trout (Salmo trutta lacustris) as biological endpoint. The egg incubation and hatching success of the salmonids and their dependence on temperature and stream substratum quality were compared. Hatching rates of Danube salmon were lower than of brown trout, probably due to higher oxygen demands and increased interstitial respiration in spring. Increases in maximum water temperature reduced hatching rates of resident and migratory brown trout (both fall‐spawners) but were positively correlated with hatching rates of Danube salmon (a spring‐spawner). Significantly longer incubation periods of resident and migratory brown trout coincided with relatively low stream substratum quality at the end of the egg incubation. Danube salmon seem to avoid low oxygen concentrations in the hyporheic zone by faster egg development favored by higher water temperatures. Consequently, the prediction of effects of temperature changes and altered stream substratum properties on gravel‐spawning fishes and biological communities should consider the observed species‐specific variances in life‐history strategies to increase conservation success.     

Design of Emergence Test Arenas Can Affect the Results of Boldness Assays

In this study, we investigate whether and how arena design of emergence tests (a commonly used boldness assay where the latency to emerge from a start box is measured) can affect the behaviour of the subject animals. We used two populations of wild brown trout fry, captured on their hatching grounds, and measured emergence latency in arenas with two different sizes of start‐box gates and two different environments into which the fish emerges from the start box (barren and complex), in a factorial design. We found that arena design affected the behaviour of the fish, but only in one of the populations. In the affected population, a large start‐box gate in combination with a barren environment reduced emergence latency. Furthermore, the time from leaving the box until entering another compartment on the opposite side of the arena was shorter in a barren environment as compared to a more complex environment. We also found that smaller brown trout fry generally showed lower tendency for emerging from the start box at all. The results of this study suggest that direct comparisons among results from experiments using different arena designs may be problematic and highlight the importance of reporting details of arena design in publications.     

The influence of habitat availability and adult density on non-diadromous galaxiid fry settlement in New Zealand

This study investigated factors controlling settlement and initial density of two species, Galaxias vulgaris and Galaxias paucispondylus (Galaxiidae), in New Zealand streams. Factors affecting maximum fry abundance were assessed at 10 sites (eight streams) differing in habitat characteristics and the presence or absence of introduced brown Salmo trutta and rainbow trout Oncorhynchus mykiss . Regression analysis indicated that conspecific adult numbers, extent of backwater habitat and brown and rainbow trout density were the most important factors determining initial fry density. Experimental additions of backwater habitat increased fry settlement tenfold compared with control reaches, demonstrating that the area available for fry settlement had a substantial effect on fry abundance. Results from this study indicate that fry settlement may be a critical limiting phase for some non-diadromous galaxiid populations. Increasing settlement area to enhance fry retention offers a practical conservation tool for fisheries managers that may aid conservation of threatened fishes like galaxiids.     

Short-and long term niche segregation and individual specialization of brown trout (Salmo trutta) in species poor Faroese lakes

Trophic niche divergence is considered to be a major process by which species coexistence is facilitated. When studying niche segregation in lake ecosystems, we tend to view the niche on a one-dimensional pelagic-littoral axis. In reality, however, the niche use may be more complex and individual fidelity to a niche may be variable both between and within populations. In order to study this complexity, relative simple systems with few species are needed. In this paper, we study how competitor presence affects the resource use of brown trout (Salmo trutta) in 11 species-poor Faroese lakes by comparing relative abundance, stable isotope ratios and diet in multiple habitats. In the presence of three-spined sticklebacks (Gasterosteus aculeatus), a higher proportion of the trout population was found in the pelagic habitat, and trout in general relied on a more pelagic diet base as compared to trout living in allopatry or in sympatry with Arctic charr (Salvelinus alpinus). Diet analyses revealed, however, that niche-segregation may be more complex than described on a one-dimensional pelagic-littoral axis. Trout from both littoral and offshore benthic habitats had in the presence of sticklebacks a less benthic diet as compared to trout living in allopatry or in sympatry with charr. Furthermore, we found individual habitat specialization between littoral/benthic and pelagic trout in deep lakes. Hence, our findings indicate that for trout populations interspecific competition can drive shifts in both habitat and niche use, but at the same time they illustrate the complexity of the ecological niche in freshwater ecosystems.     

Growth‐rate variation in brown trout in small neighbouring streams: evidence for density‐dependence?

The within- and among-population variation in individual growth rate of brown trout Salmo trutta L. was studied in five small neighbouring streams (seven isolated populations) within a distance of 70 km in east Norway. Observed growth rate was only weakly correlated with predicted maximum growth rate based on laboratory models, and there was a significant interaction with site. A generalized linear model showed that growth rate was positively correlated with temperature, but also that growth rate decreased as the summer season progressed. This might indicate either a seasonal decline in food availability or appetite, or a change in energy allocation strategy. Growth rate decreased with increasing fish age, probably as an effect of sexual maturation of older fish, and differential allocation of protein and lipid among different size-groups of brown trout. After adjusting for variation in temperature, season, and fish mass, there was still significant among-site variation in growth rate. A significant part of this variation was due to variation in brown trout density and the presence or absence of Alpine bullhead Cottus poecilopus . Growth rate decreased with increasing brown trout density, and was lower in the presence than in the absence of Alpine bullhead after correcting for variation in brown trout density. This last result may indicate the presence of interspecific competition.     

Seasonal changes in the body composition of young riverine Atlantic salmon and brown trout

The body composition of protein and fat in Atlantic salmon Salmo salar and brown trout Salmo trutta before and after winter was investigated in a temperate, small river, normally ice covered from the middle of November until the end of March. Fat, protein and specific energy declined greatly in winter but were replenished rapidly in spring. Rates of decline were slower for the smallest fish, which also had the lowest specific content of fat, protein, and energy, while the differences in absolute amounts were greatest for the largest fish. The mean specific fat content was reduced by 45–70% during winter, relative to the pre-winter period (September). Mean daily reductions in specific enegy of the larger size groups of brown trout (3·7 × 10⁻³ kJ g⁻¹ day⁻¹) were almost half of the corresponding values for the largest Atlantic salmon (6·3 × 10⁻³ kJ g⁻¹ day⁻¹) during winter. A minor reduction in protein content was found during winter, with mean reductions of 6–10% in comparison to those in September. During spring the fat content was replenished rapidly, particularly for the smallest salmon fry (a threefold increase from April to June). Fat content in the larger salmon and trout increased by about 1·8 times. Based on estimated metabolic rates, digested energy during wintertime may contribute about two-thirds of the brown trout fry's energy demand. For Atlantic salmon, the corresponding value is about 50%. The winter period put considerable stress on the young salmonids living in lotic environments, in particular for the smallest fry with the lowest energy content before winter and the largest losses during winter. This should make the fry more vulnerable to adverse abiotic and biotic factors.     

Downstream migration and critical water velocities in stream channels for fry of four salmonid species

Fry of brown trout, Atlantic salmon, brook trout and lake trout were tested for downstream migration and critical velocities with a method of stepwise increasing water velocities. Each velocity was tested for 15 min before increase to the next step. Critical velocities for fry entering the free-feeding stage, defined as the stage when the fry has resorbed its yolk sac and will have to ascend from the bottom gravel to catch food, were between 0.10 and 0.25 m s⁻¹, varying among individuals and depending on species and water temperature. Downstream displacement started at lower velocities. Lake trout had the lowest critical velocity. Temperature influenced swimming performance considerably. On average, a 7°C increase in temperature resulted in a 0.05 m s⁻¹ increase in critical velocity. The fry actively searched out the low-velocity niches in the channels. Flow-sensivity gradually decreases with fry development; when the fry had reached a length of 40–50 mm they were able to tolerate water velocities higher than 0.50 m s⁻¹.     

Habitat shifts in rainbow trout: competitive influences of brown trout

Summary We compared habitat use by rainbow trout sympatric (three streams) and allopatric (two streams) with brown trout to determine whether competition occurred between these two species in the southern Appalachian Mountains. We measured water depth, water velocity, substrate, distance to overhead vegetation, sunlight, and surface turbulence both where we collected trout and for the streams in general. This enabled us to separate the effects of habitat availability from possible competitive effects. The results provided strong evidence for asymmetrical interspecific competition. Habitat use varied significantly between allopatric and sympatric rainbow trout in 68% of the comparisons made. Portions of some differences refelected differences in habitats available in the several streams. However, for all habitat variables measured except sunlight, rainbow trout used their preferred habitats less in sympatry with brown trout than in allopatry if brown trout also preferred the same habitats. Multivariate analysis indicated that water velocity and its correlates (substrate particle size and surface turbulence) were the most critical habitat variables in the interaction between the species, cover in the form of shade and close overhead vegetation was second most important, and water depth was least important.     

The natural regulation of numbers and growth in contrasting populations of brown trout, Salmo trutta, in two Lake District streams

SUMMARY. 1. This short review summarizes a long-term investigation of brown trout in two populations that probably represent opposite extremes of life histories in this polymorphic species; Bhick Brows Beck serves as a nursery for the progeny of migratory trout (mixture of sea and estuarine trout) and Wilfin Beck is populated by resident trout. 2. Population density in Black Brows Beck was always much higher than that in Wilfin Beck, and was regulated by density-dependent survival in the early stages of the life cycle. There was no evidence for similar density-dependent regulation in Wilfin Beck; simple proportionate survival occurred with fairly constant loss-rates. Survival was reduced in both populations by summer droughts and also by spates in Wilfin Beck. 3. Black Brows trout were always larger than Wilfin Beck trout of similar age; fry size at the start of the growth period was chiefly responsible for these differences. Variations in water temperature were chiefly responsible for differences in growth rates between year-classes within each population. Food intake was not a limiting factor, except in the first winter of the life cycle and for adults over 3 years old in Wilfin Beck. Variation in individual size was inversely density-dependent in Black Brows Beck and decreased with age in Wilfin Beck, these changes being due to natural (stabilizing) selection. 4. There is strong evidence for genotypic differences between the populations. The implications of this are discussed, especially the need to conserve different populations that may contain unique genetic material, and the importance of restocking with fish reared from the indigenous population that should always contain the optimum genotypes for a particular habitat. Restocking with juveniles should be done with caution because it could lead to a decrease in both numbers and size variation when the population is regulated by density-dependent mechanisms. 5. One major objective of future work should be the development and improvement of mathematical models that can be used to predict the optimum density for trout in different populations, the maximum attainable growth rate in different habitats, and the effects on trout populations of environmental changes due to natural causes (e.g. droughts and spates) or human activities.     

Estimates of survival of stream-dwelling brown trout using

Estimated monthly apparent survival of stream-dwelling brown trout Salmo trutta in south-east Norway was higher in winter than in summer, and lower in Alpine bullhead Cottus poecilopus sites than in allopatric sites. Apparent survival denotes true survival x local site fidelity. Emigration may also explain differences in apparent survival. All brown trout included in this Study were at least 1 year old.     

Feeding pattern and diet of first feeding brown trout fry under natural conditions

The feeding habits of brown trout Salmo trutta fry were studied during the critical first feeding period in a natural spawning and nursery stream. A low proportion of the fry initiated exogenous feeding before emergence from the gravel, and while having nearly 30% of the yolk sac remaining. This probably reflected low feeding motivation or limited feeding opportunities within the gravel environment. The majority of the fry started feeding after emergence, and after most or all yolk was absorbed. Some fry emerged with large amounts of yolk remaining, while others emerged after yolk exhaustion. The degree of stomach fullness revealed that feeding was more efficient after a territory had been acquired. The diets of the young fry were dominated by chironomid larvae, followed by zooplankton and Plecoptera larvae. Fry dispersing downstream and out of the nursery area were significantly smaller than resident fry, indicating displacement due to competition for territories. The majority of the downstream dispersing fry had initiated feeding, and there was at this point no evidence of starvation in any of the fry. It therefore appeared that the later emerging fry actively migrated out of the overpopulated nursery area to find available territories further downstream.