Environmentally induced spatio-temporal variations in the fecundity of brown trout Salmo trutta L.: trade-offs between egg size and number

1. Resident brown trout Salmo trutta in the Esva River basin (north Spain) live in a patchy environment with tracts of riparian forest or meadow along stream banks. This study assessed whether the reproductive traits of brown trout from four contrasting sites reflected site-specific factors.
2. Length at maturity (10.5–11 cm of 1 + individuals) was the same in the four sites examined but slowest growers in slow-growing sub-populations delayed maturity for 1 year relative to fast-growing fish. The analysis of monthly variations in egg size and number suggest that two 'decisions' in two consecutive years are required to complete spawning. The first concerns the number of eggs, determined when trout are still 0 +, and the second concerns egg size.
3. At three sites, egg size and number did not differ significantly between years but highly significant interannual variations were apparent at another site. Fish length was the major determinant of egg size and number at all sites but for any given length, brown trout at sites where the fish exhibited higher growth rates spawned more, but smaller, eggs than those at slow-growing sites. This spatial pattern was identical to the temporal pattern exhibited by trout at another site. The combination of temporal (year-to-year) and spatial (between rivers) variations in egg size and number showed a significant negative correlation, supporting the operation of a trade-off between these two traits.
4. The trade-off between egg size and number seems to be determined by site-specific factors, with slow-growing trout at sites which are fully covered by canopy spawning fewer, but larger, eggs than fast-growers in unshaded sites.     

Comparative analysis of introgression at three marker classes: a case study in a stocked population of brown trout

Comparative analysis of protein loci, microsatellite and mtDNA markers revealed generally comparable estimates for introgression and apparent admixture rates in stocked brown trout populations at two sites in the River Doubs (Rhône dainage, Switzerland), which are 10 km apart and which belong to the same management unit. At one site, a significant deviation between mtDNA and nuclear markers could be explained by stocking of F1 hybrids originating from crosses between hatchery females and males from the local population. Substantial differences between diagnostic protein loci and protein loci having non-fixed private alleles indicated that caution must be exercised when using genetic markers not strictly diagnostic for the distinction of the populations under investigation. Congruent estimates of introgression and apparent admixture rates between diagnostic protein loci and presumed diagnostic microsatellite loci suggest that the latter can be regarded as reliable genetic markers for the estimation of introgression in Mediterranean brown trout populations stocked with trout of Atlantic origin. Significant differences in introgression and apparent admixture rates between the two sites and between age-classes of one study site were observed. Introgression is suggested to depend on environmental factors. Significantly lower introgression rates in age-class 2+ years as compared to juvenile trout might further indicate that introduced Atlantic brown trout and hybrids decrease in proportion between age-classes 1+ and 2+ years.     

First-year survival of brown trout in three Norwegian streams

Monthly survival rates during the first year of life were estimated for the 1999 cohort of stream-resident brown trout Salmo trutta in three Norwegian streams, using capture-mark-recapture methods and the Cormack–Jolly–Seber model. It was hypothesized that reduced survival would occur during the winter. For one of the study populations, the data did support seasonal variation in survival, with monthly survival rates being lower during winter than during summer (0·65 v . 0·99). For the remaining two populations, there was no evidence for seasonal variations in monthly survival rates, but monthly survival rates were significantly different (0·87 v . 0·99). No evidence was found for size-dependent winter survival. Some marked individuals emigrated from the study sites, suggesting that survival rates were underestimated and that different survival rates among populations were partly due to different emigration rates. Net immigration of brown trout was evident at all three study sites.     

Distribution of the flagellate Hexamita salmonis Moore, 1922 and the microsporidian Loma salmonae Putz, Hoffman and Dunbar, 1965 in brown trout, Salmo trutta L., and rainbow trout, Salmo gairdneri Richardson, in the River Itchen (U.K.) and three of its fish farms

The occurrence of Hexamita salmonis Moore, 1922 and Loma salmonae Putz, Hoffman and Dunbar, 1965 was investigated at 10 sites on the R. Itchen (five for brown trout only, three for rainbow trout only, and two for both brown trout and rainbow trout) and at three of its nine fish farms (two for rainbow trout, one for brown trout). Hexamita salmonis was recorded in brown trout from three river sites and the farm, and in rainbow trout from both farms and four river sites. Prevalence of Hexamita salmonis in farmed rainbow trout was higher than in farmed brown trout and was consistent with the former species being more susceptible to infection. H. salmonis was at significantly higher prevalence in rainbow trout from farm no. 5 than farm no. 2 for three size classes of fish. In wild brown trout and feral rainbow trout, the highest prevalences of H. salmonis were recorded at sites in the vicinity of farm no. 2. This distribution was consistent with an area of naturally high infection levels, and with infected fish unintentionally released from farm no. 2 serving as a source of infection, the infection subsequently becoming established in the river fish. Loma salmonae was recorded in wild brown trout and in rainbow trout from both farms. This appears to be the first recording of this parasite from British salmonids and also the first recording of the parasite from brown trout. The distribution of the parasite (particularly the prevalence being higher at farm no. 2 than farm no. 5) was consistent with it being introduced into the R. Itchen via rainbow trout from farm no. 2 (and probably no. 3) much of whose stock derived from imported Californian 'Shasta' rainbow trout.     

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.     

Temporal and spatial segregation of spawning in sympatric populations of Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

Based on data from Norwegian streams with sympatric populations of Atlantic salmon and brown trout, it is suggested that temporal segregation is the main mechanism segregating Atlantic salmon and brown trout during spawning. Peak spawning of trout was about 15 days earlier than that of salmon. Physical factors, such as water depth, water velocity and distance from the river banks segregate spawning sites of salmon and trout poorly. Gravel sizes of the redds of salmon and trout were significantly different, though with a considerable overlap, and mean egg depth of salmon and trout were 0.18 and 0.12 m, respectively, probably attributable to the different size of spawners of salmon and trout. None of the temporal or spatial parameters analysed segregate spawners of salmon and trout completely. Species determination of eggs and alevins from the redds showed no interspecific superimposition of redds. It is, therefore, concluded that low survival of hybrids after hatching does not explain the low frequency of hybrids observed in sympatric populations of salmon and trout.     

Detection and apparent survival of PIT‐tagged stream fish in winter

Environmental fluctuations exert strong control on behavior, survival, and fitness of stream biota. Technical improvements increasingly allow for tracking the response of large numbers of individuals to environmental fluctuations, for instance, by remote detection of animals equipped with PIT (passive integrated transponder) tags. PIT tags were implanted into 393 juvenile and adult brown trout Salmo trutta L. and European sculpin Cottus gobio L. in a boreal stream subjected to considerable ice formation. With weekly trackings over 6 months, we quantified apparent survival and detection probability in relation to biological, environmental, and methodological factors. Individuals with a higher physical condition in autumn showed a higher apparent survival; this pattern was consistent across all species and age classes. Detection probability decreased with increasing thickness of the surface ice layer; this effect was most pronounced for juvenile trout and benthic‐living sculpin, both tagged with smaller‐sized tags. Detection probability was reduced in structurally complex habitats. Our study demonstrates that apparent survival and particularly detection probability may show pronounced spatiotemporal variation. In order to compare results from different sampling occasions and sites, a good knowledge of the study site and of the regulating factors is crucial.     

Assessing the status of British wild brown trout, Salmo trutta, stocks: a pilot study utilizing data from game fisheries

SUMMARY. 1. This paper describes an analysis of results from a questionnaire (Giles, 1987) which requested information on the status of game fish stocks (especially wild brown trout Salmo trutta L.) from those Game Conservancy members who own or manage fisheries. Also included are data arising from fishery log book records which are used to indicate trends in brown trout catch per unit effort (CPUE) over past decades.
2. The key finding from questionnaire returns was the widespread reporting of apparent declines of wild brown trout stocks in a total of twenty-seven sites throughout Britain. The affected waters are geographically widespread from the south-west of England through southern and south-east England, south Wales, northern England and Scotland including the Western Isles. Suspected stock declines of migratory (sea) trout, salmon Salmo salar L. and grayling Thymallus thymallus L. were also reported. Where brown trout occur alone, a significantly higher proportion of questionnaire returns recorded a stock decline compared with fisheries containing both brown and rainbow trout Salmo gairdneri Richardson.
3. Trout catch records from a small selection of differing fishery types are presented and the value of such data in fish stock assessment is discussed.     

Long-term population demographics of native brook trout following manipulative reduction of an invader

Although laboratory studies have provided evidence for negative interactions between brook trout and brown trout, it is unknown how these interactions affect larger scale demographics in a natural setting. We tested the effects of invasive brown trout on brook trout demographics by removing brown trout from a sympatric population using a before–after control-impact study design. The study was conducted across a large stream network for a period of 6 years. Abundance of brook trout increased after brown trout removal primarily as a result of increased recruitment and immigration. Size structure also shifted towards larger individuals as a result of increased growth rates and a decrease in emigration of larger trout. Size at maturity and body condition did not change after brown trout removal. Adult brook trout survival increased during the post-treatment period in both the treatment and control reach. A decrease in flood intensity during the post-treatment time period may have led to increased survival. Adult survival may not be the best metric to use when assessing interactions between trout species, especially when the subordinate species has suitable areas to emigrate.     

Microsatellite DNA data point to extensive but incomplete admixture in a marble and brown trout hybridisation zone

Marble trout, endemic to the Adriatic drainage basin, is severely threatened by hybridisation with non-native brown trout. In the present study, we analysed 12 microsatellite DNA loci to assess genetic population structure and differentiation between sympatric phenotypic marble and brown trout at nine sampling sites in the upper Etsch/Adige River system. F _ST and AMOVA analyses revealed significant genetic differentiation between marble and brown trout samples. Thus, admixture between brown and marble trout appears to be incomplete. However, factorial correspondence analysis depicted marble trout, Atlantic brown trout and intermediate genotypes. Bayesian-based individual assignment tests identified indigenous marble trout at five sampling sites. In four other samples no ‘pure’ marble trout were detected. Bidirectional, first-generation hybridisation, involving both sexes of both parental species was observed. In locations where ‘pure’ marble trout still exist, post-F1 hybridisation appears to be directed towards brown trout. This has likely slowed the rate of hybridisation between the two trout species and the decline of relic marble trout populations. Based on these results, restoration management actions are proposed, such as the abandonment of brown trout stocking activities, sharper angling policies, establishment of indigenous marble trout breeding strains and the elaboration of a conservation priority list.     

Comments on the morphology of Nicsmirnovius eximius (Kiser, 1948) (Aloninae,Anomopoda, Cladocera) in Thailand,with description of its male

The density, diet and habitat use of brown trout (Salmo trutta) and Siberian sculpin (Cottus poecilopus) were studied in the subalpine River Atna in southeastern Norway in the autumn during a six year period (1986–1991). There was an inverse relationship between the density of brown trout and Siberian sculpin. Diet overlap, as indicated by the Schoener index, was high between the two species, ranging between 0.48 and 0.86. Chironomid larvae and other aquatic insects were the most common food items for both species. Brown trout also consumed substantive amounts of surface insects. Siberian sculpin typically occupied sites with finer substrates and greater water depths than brown trout, even though there was considerable overlap in habitat use between the two species. Because the two species shared similar habitats, we suggest that the potential for species interactions exists, particularly at sites where density of sculpin is high.     

Isolation and characterization of the receptor for vitellogenin from follicles of the rainbow trout,Oncorhynchus mykiss

The isolation and characterization of the receptor for vitellogenin from follicle membranes of the rainbow trout, Oncorhynchus mykiss, is described. Follicle membrane proteins subjected to SDS-polyacrylamide gel electrophoresis and subsequently to either protein staining or ligand blotting with radiolabelled vitellogenin (¹²⁵iodine-vitellogenin) demonstrated that the vitellogenin receptor has an apparent molecular mass of 200 kD (probably comprising of two 100-kD subunits) under non-reducing conditions. The vitellogenin binding sites were identified as specific receptors: binding was saturable and the binding sites were both tissue specific to follicle membranes and exhibited ligand specificity. Scatchard analyses of specific binding data revealed a single class of binding sites with a high affinity for rainbow trout vitellogenin (K _d=8.2·10^-9 mol·1^-1). Both brown trout, Salmo trutta, vitellogenin and carp, Cyprinus carpio, vitellogenin were able to displace the radiolabelled rainbow trout vitellogenin from its receptor, although they were less effective than rainbow trout vitellogenin.Abbreviations B _max maximum number of binding sites available - BSA bovine serum albumin - bt-VTG brown trout vitellogenin - c-VTG earp vitellogenin - HEPES N-2-hydroxyethylpiperazine-N-2-ethanesulfonic acid - K _d dissociatian constant - NCM nitrocellulose membranes - PMSF phenylmethylsulphonylfluoride - rt-VTG rainbow trout vitellogenin - VTG vitellogenin     

Movement of exotic plants into coastal native forests from gardens in northern New Zealand

The anti-predator behaviours of a New Zealand freshwater crayfish (Paranephrops zealandicus) to the native long-finned eel (Anguilla dieffenbachii) and the introduced brown trout (Salmo trutta) were investigated. Crayfish modified their behaviour in the presence of both trout and eels. However, a significantly greater number of defensive chela displays and swimming responses were made to eels than trout. Crayfish were able to use chemical cues from skin mucus to detect eels but not trout. Paranephrops zealandicus is able to make some appropriate defensive behavioural responses to the introduced brown trout as well as to its native predator, the long-finned eel. However, crayfish may be at greater risk from the introduced predator because of their apparent inability to detect trout using non-contact chemical cues. This may be a reflection of the different co-evolutionary histories crayfish have had with trout and eels.     

Complex genetic diversity patterns of cryptic,sympatric brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) populations in tiny mountain lakes

Intraspecific genetic variation can have similar effects as species diversity on ecosystem function; understanding such variation is important, particularly for ecological key species. The brown trout plays central roles in many northern freshwater ecosystems, and several cases of sympatric brown trout populations have been detected in freshwater lakes based on apparent morphological differences. In some rare cases, sympatric, genetically distinct populations lacking visible phenotypic differences have been detected based on genetic data alone. Detecting such “cryptic” sympatric populations without prior grouping of individuals based on phenotypic characteristics is more difficult statistically, though. The aim of the present study is to delineate the spatial connectivity of two cryptic, sympatric genetic clusters of brown trout discovered in two interconnected, tiny subarctic Swedish lakes. The structures were detected using allozyme markers, and have been monitored over time. Here, we confirm their existence for almost three decades and report that these cryptic, sympatric populations exhibit very different connectivity patterns to brown trout of nearby lakes. One of the clusters is relatively isolated while the other one shows high genetic similarity to downstream populations. There are indications of different spawning sites as reflected in genetic structuring among parr from different creeks. We used >3000 SNPs on a subsample and find that the SNPs largely confirm the allozyme pattern but give considerably lower F _ST values, and potentially indicate further structuring within populations. This type of complex genetic substructuring over microgeographical scales might be more common than anticipated and needs to be considered in conservation management.     

Effects of pulsed discharges from a hydropower station on summer diel feeding activity and diet of brown trout (Salmo trutta Linnaeus, 1758) in an Iberian stream

The influence of pulsed discharges associated with hydroelectric power generation (i.e. hydropeaking) on feeding activity and diet composition of adult brown trout (Salmo trutta) was studied during the summer by comparing two sites: upstream (control site) and downstream from a power plant (hydropeaking site). Twenty fish were captured from each study site by electrofishing at 4‐hour intervals for two consecutive days and stomach contents were collected with pulsed gastric lavage. Hydropeaking events affected brown trout feeding behaviour as well as prey availability. Feeding intensity, measured by the stomach Fullness Index, showed pronounced variations with maximum values after flow pulses, which were linked to variations in prey availability because of increased drift rates of invertebrates. In contrast, brown trout living at the control site showed smoother variations in feeding activity not linked to invertebrate drift. Overall, brown trout at the hydropeaking site had higher food consumption rates and a more generalist and heterogeneous diet than trout from the control site, indicating an opportunistic feeding behaviour during flow pulses. Therefore, the hydrological disturbance caused by hydropeaking did not appear to cause direct negative impacts on feeding of adult brown trout. However, reduced trout density and imbalanced size structure in the hydropeaking site were detected, requiring further research to clarify the spatial influence of hydropeaking on other factors that could negatively affect brown trout populations.     

Evidence for selective angling of introduced trout and their hybrids in a stocked brown trout population

Comparisons of the genetic composition of brown trout Salmo trutta captured by anglers and by electrofishing based on three diagnostic microsatellite loci provided strong evidence that angling is selective in a stocked brown trout population. At two sites, anglers caught significantly younger trout and proportionally more introduced hatchery trout and hybrids than were observed in electrofishing surveys. Selective angling, in combination with a small legal catch size, may have considerably eliminated introduced trout and hybrids before spawning at the study sites, and thus may have reduced the introgression of alien genes into the local gene pool. Angling can be an important factor influencing the genetic structure of fish populations and should be taken into account in studies of introgressive hybridization in stocked fish populations and their management. In this study, demographic consequences of stocking were not assessed. Thus, even though the genetic consequences of stocking may be minimal or largely reversible through angling, resource competition between native and introduced trout, until they reach legal catch size, is expected to have a negative effect on the productivity of the indigenous trout population.     

A Bioenergetic Analysis of Factors Limiting Brown Trout Growth in an Ozark Tailwater River

We examined prey utilization and energy consumption by brown trout, Salmo trutta, in a cold tailwater (Little Red River, Arkansas, USA; LRR) having low biodiversity and low availability of fish as prey. Stomach content analysis and age estimation were performed on thirty brown trout (10 each of three size classes for a total of 710 trout) collected monthly from an upstream and downstream site over a 1-year period. Diet diversity was low at both sites, as 80% and 70% of all prey consumed by upstream and downstream brown trout, respectively, were isopods. Piscivory (<0.5% of individuals sampled) and consumption of terrestrial invertebrates were rare. There was no relation between diet diversity and trout age, and a very small ontogenetic shift in brown trout diet. Second, we investigated brown trout growth rates relative to prey consumption and temperature. Temperatures and availability of prey were less than required for maximal trout growth. However, prey availability limited trout growth directly, but sub-optimal temperatures probably buffered the effect of this reduced energy consumption by reducing metabolic energy expenditures. Brown trout growth was 54.8–57.0% of the maximum predicted by a bioenergetics model. Instantaneous growth rates for age 1 and adult brown trout were slightly higher for those downstream (0.195) versus those upstream (0.152). Although isopods are abundant within this tailwater to serve as a forage base, the displacement of native fish fauna and subsequent lack of establishment of cold-tolerant forage fish species due to the thermal regime of hypolimnetic release from Greers Ferry Reservoir probably serves as a major barrier to brown trout growth.     

Influence of hydrologic attributes on brown trout recruitment in low-latitude range margins

Factors controlling brown trout Salmo trutta recruitment in Mediterranean areas are largely unknown, despite the relevance this may have for fisheries management. The effect of hydrological variability on survival of young brown trout was studied during seven consecutive years in five resident populations from the southern range of the species distribution. Recruit density at the end of summer varied markedly among year-classes and rivers during the study period. Previous work showed that egg density the previous fall did not account for more than 50% of the observed variation in recruitment density. Thus, we expected that climatic patterns, as determinants of discharge and water temperature, would play a role in the control of young trout abundance. We tested this by analyzing the effects of flow variation and predictability on young trout survival during the spawning to emergence and the summer drought periods. Both hatching and emergence times and length of hatching and emergence periods were similar between years within each river but varied considerably among populations, due to differences in water temperature. Interannual variation in flow attributes during spawning to emergence and summer drought affected juvenile survival in all populations, once the effect of endogenous factors was removed. Survival rate was significantly related to the timing, magnitude and duration of extreme water conditions, and to the rate of change in discharge during hatching and emergence times in most rivers. The magnitude and duration of low flows during summer drought appeared to be a critical factor for survival of young trout. Our findings suggest that density-independent factors, i.e., hydrological variability, play a central role in the population dynamics of brown trout in populations from low-latitude range margins. Reported effects of hydrologic attributes on trout survival are likely to be increasingly important if, as predicted, climate change leads to greater extremes and variability of flow regimes.     

Changes in haematocrit values in blood samples treated with and without oxygen: a comparative study with four salmonid species

Changes in haematocrit values under lowered oxygen tension and aerobically treated blood samples of rainbow trout Salmo gairdneri (Richardson), landlocked Baltic salmon S. salar (L.), brown trout S. trutta lacustris (L.) and lake trout Salvelinus namaycush (Walbaum) have been studied in vitro . The mean haematocrit value increased during 2 h incubation under lowered oxygen tension by 32.7 ± 3.1% in rainbow trout, 28.2 ± 2.8% in landlocked Baltic salmon, 29.2 ± 5.6% in brown trout and 25.2 ± 2.8% in lake trout. During corresponding incubation with oxygen the mean haematocrit value decreased below the starting level by 18.1 ± 1.9% in rainbow trout, 18.3 ± 4.8% in landlocked Baltic salmon, 22.4 ± 0.7% in brown trout and 11.8 ± 1.7% in lake trout. Consequently, the changes in the haematocrit values were greater in the species belonging to the genus Salmo than in that of the genus Salvelinus . The increase in the haematocrit values seems to have resulted from swelling of the erythrocytes and their decrease is similarly attributable to shrinking of the cells. The reasons for this swelling, which may be complicated, and its apparent significance for haematocrit determinations are discussed.     

Caddis larvae cases (Trichoptera,Limnephilidae) as anti-predatory devices against brown trout and sculpin

Brown trout (Salmo trutta L.) and sculpin (Cottus gobio L.) predation on the cased limnephilid larvae Glyphotaelius pellucidus (Retz.) (weak, broad leaf case), Limnephilus pantodapus McLachl. (weak, long cylindrical leaf case), L. rhombicus (L.) (rigid, hedgehog case) and Potamophylax cingulatus (Steph.) (rigid, cylindrical mineral case) was studied in the laboratory. The proportion of larvae ingested was significantly higher in brown trout than in sculpin for all four prey species. Brown trout captured larvae of all four species with equal success, but the higher number of unsuccessful captures resulted in a higher survival rate for P. cingulatus. It was significantly more difficult for sculpin than for brown trout to capture L. pantodapus larvae and to ingest all species except L. rhombicus. P. cingulatus larvae were never ingested by sculpins, and had the highest survival rate after capture by both fish species. In brown trout, the handling time of L. pantodapus was significantly longer than that of P. cingulatus. Sculpin had a significantly longer handling time of G. pellucidus than brown trout. L. pantodapus and L. rhombicus larvae feigned dead significantly longer than did those of G. pellucidus and P. cingulatus upon attacks from sculpins. L. rhombicus larvae also feigned dead for a long time upon attacks from brown trout. The survival rate of a larva attacked by sculpin or brown trout depends on both case rigidity as well as its behaviour. Brown trout was a more successful predator of cased caddis larvae than sculpin.