Predation by brown trout: a major mortality factor for sexually mature European minnows

Brown trout Salmo trutta in the subalpine lake, Øvre Heimdalsvatn, showed large temporal variation in the rate of predation on the introduced minnow Phoxinus phoxinus population. Minnows were found in the stomachs of brown trout between 16 and 38 cm L _T. Significantly greater predation was recorded shortly after ice break at the end of June 1999, with frequencies of 9 and 20% within the L _T classes 16–29·9 cm and  ≥30 cm, respectively. Predation on minnows was only occasionally detected during July, August and September. The high level of predation coincided with minnow spawning, and lengths of consumed minnows were equal to those of sexually mature individuals. Accepting a causal link between minnow spawning, which lasted c . 3 weeks, and the contemporary high rate of predation, the estimated annual consumption of minnows by the brown trout population would be 138 kg wet mass. Although most of the annual consumption of minnows by brown trout (90%) occurred within a very short period (3 weeks), it accounted for a significant proportion (60%) of the annual loss in biomass of the sexually mature part of the population.     

The brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in the lake, Øvre Heimdalsvatn: long-term changes in population dynamics due to exploitation and the invasive species,European minnow (<Emphasis Type="Italic">Phoxinus phoxinus</Emphasis>)

Studies of the brown trout (Salmo trutta) population in the Norwegian subalpine lake, Øvre Heimdalsvatn, over a 50-year period have revealed major changes in population dynamics. In 1958, the population density was high, with individuals stagnating in growth at lengths below 30 cm. After heavy exploitation during the years 1958–1969, the number of older fish declined substantially, and growth rates increased significantly. Since 1969, the European minnow (Phoxinus phoxinus) have been observed in the lake, with increasing densities from 1977–1978 to 1999–2000. The age structure of the brown trout population has changed markedly from the 1960s to the period 1993–2006. Annual recruitment significantly declined, from an average number of 3746 individuals in age-class 4 during the period 1958–1966 to an average of 1668 during the period 1993–2006. However, due to lower exploitation rates, the number of old fish was significantly higher in the latter period. The summer diet of brown trout has changed substantially from a dominance of the large crustaceans Lepidurus arcticus and Gammarus lacustris to a high occurrence of European minnows, while L. arcticus has become practically absent from the diet. There was a negative relationship between brown trout biomass and annual length increment. However, despite a brown trout biomass at the same level during the years 1993–2006 as in the 1960s, annual individual growth rates have significantly declined. The reduced recruitment and reduced annual growth rates of the brown trout, as well as changes in the diet, are most likely related to the introduction and establishment of the invasive species, the European minnow.     

Species-specific effects of gastropods on leaf litter processing in pond mesocosms

A study of the diet of native brown trout (Salmo trutta) parr and introduced European minnow (Phoxinus phoxinus) in the subalpine lake, Øvre Heimdalsvatn, showed that the two species had considerable dietary overlap, both in the littoral zone and in the outlet of the lake. Chironomidae constituted a substantial proportion of the diet of the two species in both habitats. The results indicated that both zooplankton (Cladocera) and large macroinvertebrates (EPT-species) made up a higher proportion of the minnow diet in the early phase of the minnow establishment (1975–1977) than later, and that the significance of small macroinvertebrates (Chironomidae) as prey has increased during the same period. Dietary analysis of the sympatric brown trout and minnow population in Øvre Heimdalsvatn does not provide a definitive conclusion about the degree of competition between the two species. However, together with the findings in other studies in Øvre Heimdalsvatn that have documented reduced recruitment and individual growth of brown trout, decreased individual growth of minnows and a marked decline in the density of large crustaceans (Lepidurus arcticus and Gammarus lacustris) in the shallow littoral of the lake during the last decades indicates that competitive interactions between the two species are likely. This is probably an example of the competition between two fish species with a high degree of dietary overlap when living in sympatry, most likely caused by absence of alternative prey and alternative habitats due to the high predation risk for both brown trout parr and minnows in deeper parts and in the open waters of the lake.     

Biology of the minnow Phoxinus phoxinus and its interactions with brown trout Salmo trutta in Øvre Heimdalsvatn, Norway

Minnows have recently been introduced into the mountain lakes of southern Norway. In Øvre Heimdalsvatn, an intensively studied lake, the minnow population has increased rapidly during the last decade. Studies were performed on feeding habits, habitat preferences, age, growth, energy content, densities, parasites, predation and spawning activities. Several aspects were found to be different from other studies, e.g. higher age of sexual maturation, lower growth rate, no feeding on plants, and low infection of parasites. The interaction with the only other fish, the brown trout, is discussed in relation to the minnow population. In 1978 the minnows constituted little more than 10% of production and food consumption of the trout population, and only limited food competition and predation was documented. Increasing competition is likely with a greater minnow population.     

The history of the minnow Phoxinus phoxinus (L.) in Norway: from harmless species to pest

The causes, effects and extent of minnow Phoxinus phoxinus introductions in Norway are reviewed to assess why the introductions have had severe effects, especially where brown trout Salmo trutta is the only fish species present. The natural distribution of minnow in Norway was mainly restricted to low altitude localities in the south-eastern part of the country and in some northern areas. The distribution area expanded considerably throughout the 1900s, especially in mountain areas, due in part to the use of minnows as live bait for angling. Although minnow densities do not seem unusually high in the relatively complex fish communities of its native range, the species can achieve very high population densities when introduced to communities with few fish species, such as in the numerous recently invaded lakes where brown trout was the only fish species present. The dense minnow populations in these lakes appear to have led to reduced recruitment and growth rates in the brown trout, with abundances on average 35% lower in lakes where minnow has been introduced. The success of minnow in harsh habitats demonstrates their phenotypic and ecological plasticity, but also implies that their original distribution in Norway was restricted by early immigration history and not by environmental limitations. This suggests that human-assisted spread of the species could have strong adverse effects in Scandinavia lakes of low fish species richness.     

Histopathological and ultrastructural observations of metacercarial infections of Diplostomum phoxini (Digenea) in the brain of minnows Phoxinus phoxinus

The spatial distribution and histopathological changes induced by metacercariae of the digenean trematode Diplostomum phoxini (Faust, 1918) in the brains of European minnows Phoxinus phoxinus (L.) from the River Endrick, Scotland, were studied by light and electron microscopy. Post-mortem examination of a sample of 34 minnows revealed that 50% (n = 17) of the population was infected with 13.7 +/- 2.6 (mean +/- SE; range 1 to 38) metacercariae per infected host. Serial histological sections of the infected minnow brains revealed that the metacercariae were unevenly distributed throughout the brain, with aggregations occurring in the cerebellum, the medulla oblongata and the optic lobes. In fish with highest intensities of infection, over 40% of the cerebellar area and about 30% of the medulla oblongata area were occupied by larvae. Metacercariae disrupt the integrity of brain tissue, with individuals being found in small pockets surrounded by cellular debris. Metacercariae were rarely encountered on the surface of the brain. Electron microscopic examination of infection sites revealed that the granular layer surrounding metacercariae was necrotic, exhibited nuclear degradation and was marked by vacuolation of the cytoplasm. Rodlet cells, the only inflammatory cell types recorded in this study, were found only in parasitized brains and in close proximity to the teguments of metacercariae. It is hypothesised that secretions released from the teguments of metacercariae are a counter response to protect the metacercariae from the fish brain's cellular defence mechanisms.     

Potential for parasite-induced biases in aquatic invertebrate population studies

Recent studies highlight the need to include estimates of detection/capture probability in population studies. This need is particularly important in studies where detection and/or capture probability is influenced by parasite-induced behavioral alterations. We assessed potential biases associated with sampling a population of the amphipod Gammarus lacustris in the presence of Polymorphus spp. acanthocephalan parasites shown to increase positive phototaxis in their amphipod hosts. We trapped G. lacustris at two water depths (benthic and surface) and compared number of captures and number of parasitized individuals at each depth. While we captured the greatest number of G. lacustris individuals in benthic traps, parasitized individuals were captured most often in surface traps. These results reflect the phototaxic movement of infected individuals from benthic locations to sunlit surface waters. We then explored the influence of varying infection rates on a simulated population held at a constant level of abundance. Simulations resulted in increasingly biased abundance estimates as infection rates increased. Our results highlight the need to consider parasite-induced biases when quantifying detection and/or capture probability in studies of aquatic invertebrate populations.     

伊比利亚河共生的当年生鳟和埃布罗河鳟的夏季摄食关系

1996-1998年8月在Larraun河(埃布罗河流域,西班牙北部)共捕获306尾埃布罗河(鱼岁)和185尾0龄的鳟,分析比较了胃含物组成,并测定了种间的食物重叠.食物的个数百分比组成表明两种都主要摄食水生无脊椎动物.摄食策略图显示两种都是摄食不同食物 种类的广食性种类.另外,比较胃含物和底栖大型无脊椎动物群落表明,0龄的鳟和埃布罗河(鱼岁)不摄食光螺科、钩虾科、细蜉科、 四节蜉科和卷襀科的物种,而喜食摇蚊科、五节蜉科和毛翅目物种.尽管简化的Morisitas指数表明Larraun河中这两种鱼之间的食物重叠是显著的,但由于栖息地不同和摄食的可塑性可以降低种间的竞争,使得该水域的这两种鱼能以相对较大的数量共存.然而,在食物资源和可用栖息地有限的河域,埃布罗河(鱼岁)的引入,其适应性和可塑性有可能威胁单独存在的鳟     

Comparative susceptibility of rainbow trout Oncorhynchus mykiss and brown trout Salmo trutta to Myxobolus cerebralis, the cause of salmonid whirling disease.

The susceptibility of rainbow trout Oncorhynchus mykiss and brown trout Salmo trutta to Myxobolus cerebralis, the cause of salmonid whirling disease, was assessed following dosed exposures to the infectious stages (triactinomyxons). Parallel groups of age-matched brown trout and rainbow trout were exposed to 10, 100, 1000 or 10,000 triactinomyxons per fish for 2 h and then placed in aquaria receiving single pass 15 degrees C well water. Severity of infection was evaluated by presence of clinical signs (whirling and/or black tail), prevalence of infection, severity of microscopic lesions, and spore counts 5 mo after exposure. Clinical signs of whirling disease, including a darkened caudal region (black tail) and radical tail chasing swimming (whirling), occurred first among rainbow trout at the highest dose at 6 to 7 wk post exposure. Black tail and whirling occurred among rainbow trout receiving 1000 and 100 triactinomyxons per fish at 8 to 9 wk post exposure. Only 1 of 20 fish had a black tail among rainbow trout receiving 10 triactinomyxons per fish, although 30% of the fish were infected at 5 mo post exposure. Black tails were observed in brown trout at 1000 and 10,000 triactinomyxons per fish beginning at 11 and 7 wk post exposure, respectively. There was no evidence of the tail chasing swimming (whirling) in any group of brown trout. The prevalence of infection, spore numbers, and severity of microscopic lesions due to M. cerebralis among brown trout were less at each exposure dose when compared to rainbow trout. Infections were found among rainbow trout at all doses of exposure but only among brown trout exposed to doses of 100 triactinomyxons per fish or greater. Risk of infection analyses showed that rainbow trout were more apt to be infected at each exposure dose than brown trout. Spore counts reached 1.7 x 10(6) per head among rainbow trout at the highest dose of exposure compared to 1.7 x 10(4) at the same exposure dose among brown trout. Spore numbers increased with dose of exposure in rainbow trout but not in brown trout. As microscopic lesion scores increased from mild to moderate, spore numbers increased in rainbow trout but not brown trout. The mechanisms by which brown trout resist infections with M. cerebralis were not determined. Cellular immune functions, including those of eosinophilic granular leukocytes that were more prominent in brown trout than rainbow trout, may be involved.     

Determination of factors influencing stomach content retention by striped bass captured using gillnets

The influence of mode of capture, season of capture and total body length ( L _T) on the probability of regurgitation for striped bass Morone saxatilis captured using gillnets in Smith Mountain Lake, Virginia, was examined. Overall, the mean rate of regurgitation for striped bass which contained food contents in their stomach was 8·3%. Striped bass captured by wedging had a higher mean regurgitation rate (17%) than individuals that were either entangled (5%) or gilled (2%). Striped bass caught during the autumn had approximately the same frequency of regurgitation as individuals captured during the summer (10 v . 9%), but these regurgitation rates were higher than those observed for fish during the spring sampling periods (4%). Larger striped bass were more likely to regurgitate their stomach contents than smaller individuals, with the frequency of regurgitation increasing by 0·7% for every 50 mm increase in L _T. The results of this study demonstrate the importance of identifying factors that influence regurgitation of stomach contents to minimize and account for biases associated with diet data collected from striped bass captured in gillnets. Using this information, sampling recommendations for food‐habit and feeding‐rate studies involving the collection of piscivorous fishes using gillnets are made.     

Can prey exhibit threat-sensitive generalization of predator recognition? Extending the Predator Recognition Continuum Hypothesis

Despite the importance of predator recognition in mediating predator-prey interactions, we know little about the specific characteristics that prey use to distinguish predators from non-predators. Recent experiments indicate that some prey who do not innately recognize specific predators as threats have the ability to display antipredator responses upon their first encounter with those predators if they are similar to predators that the prey has recently learned to recognize. The purpose of our present experiment is to test whether this generalization of predator recognition is dependent on the level of risk associated with the known predator. We conditioned fathead minnows to chemically recognize brown trout either as a high or low threat and then tested the minnows for their responses to brown trout, rainbow trout (closely related predator) or yellow perch (distantly related predator). When the brown trout represents a high-risk predator, minnows show an antipredator response to the odour of brown trout and rainbow trout but not to yellow perch. However, when the brown trout represents a low-risk predator, minnows display antipredator responses to brown trout, but not to the rainbow trout or yellow perch. We discuss these results in the context of the Predator Recognition Continuum Hypothesis.     

Effect of Pomphorhynchus laevis (Acanthocephala) on putative neuromodulators in the intestine of naturally infected Salmo trutta

Immunohistochemical and pathological studies were carried out on the digestive tract of parasitized and uninfected specimens of Salmo trutta (L.). A total of 124 brown trout were collected on several occasions from 3 tributaries of the Brenta River, northern Italy. Twenty-eight individuals of S. trutta (22.6%) were parasitized with Pomphorhynchus laevis (Miller, 1776). The occurrence of P. laevis in the trout gut significantly increased the number of endocrine cells immunoreactive to calcitonin gene-related peptide (CGRP), beta-endorphin, met-enkephalin, neuropeptide Y (NPY) and Substance P (SP) antisera. Moreover, bombesin-, cholecistokinin-8- (CCK-8), leu-enkephalin- and serotonin- (5-HT)-like immunoreactive cells were less numerous in the intestine of the parasitized brown trout. A strong positive immunoreactivity was observed in nerve fibres and neurones of the myenteric plexus of the parasitized fish; the antisera involved in this positive reactivity were bombesin, met-enkephalin, SP and vasoactive intestinal peptide (VIP). More neurones immunoreactive to anti-CGRP and anti-5-HT sera were noted in the myenteric plexus and in the inner layer of the tunica muscularis of the infected fish. Most of the above-mentioned neuromodulators are known to control gut motility, digestive/absorptive processes, as well as the immune response. The changes induced by parasites in the neuroendocrine system of the brown trout are discussed.     

Rainbow trout (Salmo gairdneri) stocking and Contracaecum spp

A stocking program with rainbow trout (Salmo gairdneri) at High Rock Lake, Manitoba failed due to infections with large numbers of Contracaecum spp. larvae. Nematode larvae in the intestinal tract, body cavity and musculature made the fish unmarketable. A combination of experimental infections of rainbow trout and pelicans (Pelecanus erythrorhynchos), observations on the behavior of fish-eating birds, and numbers of larval Contracaecum spp. in minnow species led to the following conclusions. The introduction of rainbow trout attracted large numbers of fish-eating birds, particularly pelicans. Concurrent predation by rainbow trout on fathead minnows (Pimephales promelas), five-spined sticklebacks (Culaea inconstans), and nine-spined sticklebacks (Pungitius pungitius), concentrated the parasites. The combined increase in densities of the introduced fish host and fish-eating birds, and the short life cycle of the parasite, increased the numbers of parasites in rainbow trout over a season and in the indigenous minnow species between years. Numbers of larvae in the indigenous minnow species declined when stocking of rainbow trout was stopped and use of the lake by fish-eating birds, particularly pelicans, returned to normal levels.     

Detection of VHSV IVb within the gonads of Great Lakes fish using in situ hybridization

Viral haemorrhagic septicaemia virus (VHSV) genotype IVb was recently detected as the cause of numerous mortality events in Great Lakes fish. In situ hybridization was used to examine the gonads from 13 fish, including freshwater drum Aplodinotus grunniens and muskellunge Esox masquinongy that were infected naturally, as well as rainbow trout Oncorhynchus mykiss and fathead minnows Pimphales promelas, which were experimentally infected. Although the ovaries and testes of fish infected by VHSV IVb had few lesions, viral RNA was present in the ovaries of the rainbow trout and fathead minnow and was abundant in the gonads of muskellunge and in the ovaries of freshwater drum. Viral RNA was present mainly surrounding yolk vacuoles/granules or adjacent to the germinal vesicle, with lesser amounts found within the germinal vesicle, in the mesovarium and/or tunica albuginea and blood vessels of the ovary. Viral RNA was also found in and surrounding primary and secondary spermatocytes of the muskellunge.     

Mortality of Baltic sea trout (Salmo trutta) after release from gillnets

We explore the mortality rate of disentangled sea trout in whitefish fishery using gillnets with a 35–43 mm bar length. The study was conducted during the main fishing seasons in the Gulf of Bothnia in the northern Baltic Sea. Overall 59.5% of the sea trout were alive at the end of a 2–7 day observation period following release from the gillnets. Altogether, 12.1% of the captured fish were found dead in gillnets and 28.4% died due to injuries during an extended observation period. The average length of the captured sea trout was 435 mm, indicating that the majority were spending their first or second year at sea. The proportion of the survived and not injured (no observed damage) fish was highest in larger fish, >450 mm. The injured and not injured fish died equally frequently. Post capture survival was not connected to the removal time from gillnets or type of observed injuries but to the type of entanglement. Most of the fish were entangled by a mesh around the body, which caused extensive scale loss and open sores on the skin. The smallest fish may have had internal wounds that were not registered in this study. These results can be used in fisheries management to estimate the mortality of multi‐species gillnet fishing to sea trout populations in relation to management actions.     

Plasticity of diel patterns in the diet and habitat use of feral, non-native fathead minnow Pimephales promelas (Actinopterygii, Cyprinidae) in a pond-dwelling population in England

The aim of the study was to test for diel patterns in the diet and habitat use in a feral, pond-dwelling, non-native fathead minnow Pimephales promelas population in England. Fish were collected in June 2009 using traps set in four habitat types (open waters, rushes??Juncus effusus, weeds??Potamogeton natans and mixed vegetation), then sexed and measured for total length and eviscerated weight in the laboratory. Data were analysed at 6-h intervals, with Fulton??s condition index and three dietary parameters (frequency of occurrence, number and weight) calculated. Minor diurnal differences in habitat use were observed in males, females and immatures, and these may be due to predator avoidance. Body condition varied greatly in rushes during daytime, probably due to shifts in habitat suitability (e.g. food, refuge). Detritus dominates the diet of native fathead minnow; however, planktonic crustaceans were the most important food resource for this population, with a clear ontogenetic shift, irrespective of habitat, towards greater proportion of ingested detritus in larger individuals. Overall, the results demonstrate that feral fathead minnows display substantial trophic plasticity and a wide range of habitat use, which is normally associated with invasive species. However, established fathead minnow populations in Europe are rare despite its wide-spread ornamental and scientific use.     

Fish-Community Response To Protective Liming in Thrush Lake, Minnesota

A protective limestone treatment was applied to an acid-sensitive lake in northeastern Minnesota as part of the Acid Precipitation Mitigation Program. This 6–year study evaluated the impact of that treatment on lakes in the upper Midwest that experience episodes of acid stress but have not lost basic species integrity and community structure. Several changes in the fish community can be directly or indirectly attributed to the addition of 4.6 tonnes of calcium carbonate early in the third year of the study. An almost 30–fold increase in the population of Pimephales promelas(fathead minnow) a year after liming, based on mark-recapture estimates from trap netting and snorkeling, was attributed to a pH increase and a three-fold increase in the calcium concentration of the epilimnion. After the initial increase, the abundance of fathead minnows declined in subsequent years, as did the elevated pH and calcium concentrations. The Salvelimis fontinalis(brook trout) population also increased in the lake following application of limestone, but this was due in part to closing the lake to fishing. An increase in survival of stocked brook trout to age 1+ and an increase in growth of older brook trout after liming were attributed to the increased forage that the fathead minnows provided. Fathead minnows may have also reduced predation pressure on young brook trout by older brook trout. This study demonstrated that liming of a slightly acidic lake did not adversely affect the integrity of the fish community, and in fact may have increased the abundance and biomass of the forage fish community and indirectly increased the survival, abundance, and growth of brook trout.     

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.     

The role of rodlet cells in the inflammatory response in Phoxinus phoxinus brains infected with Diplostomum

European minnows, Phoxinus phoxinus L., are commonly infected with Diplostomum phoxini Faust, 1918 metacercariae. A sub-sample of 34 minnows collected from the River Endrick, Stirlingshire, Scotland revealed that 50% of the population were infected (n = 17), with the cerebellum, medulla oblongata and the optic lobe regions of the minnow brain bearing the heaviest infections (13.7 +/- 2.6 mean +/- S.E.; 1-38 range). Serial histological sections through the brains of both uninfected and infected minnows revealed the presence of rodlet cells in the latter, which were occasionally observed in close proximity to the tegument of a metacercaria. Rodlet cells were the only type of host inflammatory cells found in this study and their role in the host's immune response to parasitic infection is commented upon.     

伊比利亚河共生的当年生鳟和埃布罗河的夏季摄食关系(英文)

1996-1998年8月在Larraun河(埃布罗河流域,西班牙北部)共捕获306尾埃布罗河和185尾0龄的鳟,分析比较了胃含物组成,并测定了种间的食物重叠。食物的个数百分比组成表明两种都主要摄食水生无脊椎动物。摄食策略图显示两种都是摄食不同食物种类的广食性种类。另外,比较胃含物和底栖大型无脊椎动物群落表明,0龄的鳟和埃布罗河不摄食光螺科、钩虾科、细蜉科、四节蜉科和卷襀科的物种,而喜食摇蚊科、五节蜉科和毛翅目物种。尽管简化的Morisitas指数表明Larraun河中这两种鱼之间的食物重叠是显著的,但由于栖息地不同和摄食的可塑性可以降低种间的竞争,使得该水域的这两种鱼能以相对较大的数量共存。然而,在食物资源和可用栖息地有限的河域,埃布罗河的引入,其适应性和可塑性有可能威胁单独存在的鳟。