Stocked trout have minimal effects on littoral invertebrate assemblages of productive fish‐bearing lakes: a whole‐lake BACI study

1. Rainbow Trout (Oncorhynchus mykiss [Walbaum]) is commonly stocked as a sport fish throughout the world but can have serious negative effects on native species, especially in headwater systems. Productive fish‐bearing lakes represent a frequently stocked yet infrequently studied system, and effects of trout in these systems may differ from those in headwater lakes. 2. We used a Before‐After Control‐Impact (BACI) design to determine how stocked trout affected assemblage‐level and taxon‐level biomass, abundance and average length of littoral invertebrates in a stocked lake relative to three unstocked control lakes in the boreal foothills of Alberta, Canada. Lakes were studied 1 year before and for 2 years after stocking. Because characteristics of productive fish‐bearing lakes should buffer impacts of introduced fish, we predicted that trout would not affect assemblage‐level structure of littoral invertebrates but might reduce the abundance or average length of large‐bodied taxa frequently consumed by trout. 3. Relative to the unstocked control lakes, biomass, but not abundance, of the littoral invertebrate assemblage was affected indirectly by trout through increases of some taxa after trout stocking. At the individual taxon‐level, trout stocking did not affect most (23 of the 27) taxa, with four taxa increasing in abundance or biomass after stocking. Only one taxon, Chironomidae, showed evidence of size‐selective predation by trout, being consumed frequently by trout and decreasing significantly in average length after stocking. 4. Our results contrast with the strong negative effects of trout stocking on invertebrate assemblages commonly reported from headwater lakes. A combination of factors, including large and robust native populations of forage fish, the generalised diet of trout, overwinter aeration, relatively high productivity and dense macrophyte beds, likely works in concert to reduce potentially negative effects of stocked trout in these systems. As such, productive, fish‐bearing lakes may represent a suitable system for trout stocking, especially where native sport fish populations are lacking.     

Geography of Invasion in Mountain Streams: Consequences of Headwater Lake Fish Introductions

The introduction of fish into high-elevation lakes can provide a geographic and demographic boost to their invasion of stream networks, thereby further endangering the native stream fauna. Increasingly, remaining populations of native salmonids are concentrated in fragmented headwater refugia that are protected by physical or biological barriers from introduced fishes that originate in the pervasive source populations established at lower elevations. Although fish introduced near mainstem rivers frequently encounter obstacles to upstream dispersal, such as steep slopes or falls, we found that brook trout (Salvelinus fontinalis) dispersed downstream through channel slopes of 80% and 18-m-high falls. Thus, headwater lake stocking provides source populations that may be capable of invading most downstream habitats, including headwater refugia of native fishes. The extent of additional area invasible from lakes, beyond that invasible from downstream, depends on the geography of the stream network, particularly the density and distribution of headwater lakes and their location relative to barriers inhibiting upstream dispersal. In the thermal and trophic environments downstream of lakes, fish commonly grow faster and thus mature earlier and have higher fecundity-at-age than their counterparts in other high-elevation streams. The resulting higher rates of population growth facilitate invasion. Larger body sizes also potentially aid the fish in overcoming barriers to invasion. Trout introductions to high-elevation headwater lakes thus pose disproportionately large risks to native fishes—even when the place of introduction may appear to be spatially dissociated from populations of the native species. Mapping the potential invasible area can help to establish priorities in stocking and eradication efforts. Received 28 March 2000; accepted 9 February 2001.     

Loss of genetic integrity correlates with stocking intensity in brook charr (Salvelinus fontinalis)

Supportive breeding and stocking performed with non‐native or domesticated fish to support sport fishery industry is a common practice throughout the world. Such practices are likely to modify the genetic integrity of natural populations depending on the extent of genetic differences between domesticated and wild fish and on the intensity of stocking. The purpose of this study is to assess the effects of variable stocking intensities on patterns of genetic diversity and population differentiation among nearly 2000 brook charr (Salvelinus fontinalis) from 24 lakes located in two wildlife reserves in Québec, Canada. Our results indicated that the level of genetic diversity was increased in more intensively stocked lakes, mainly due to the introduction of new alleles of domestic origin. As a consequence, the population genetic structure was strongly homogenized by intense stocking. Heavily stocked lakes presented higher admixture levels and lower levels of among lakes genetic differentiation than moderately and un‐stocked lakes. Moreover, the number of stocking events explained the observed pattern of population genetic structure as much as hydrographical connections among lakes in each reserve. We discuss the implications for the conservation of exploited fish populations and the management of stocking practices.     

The impact of stocking on the genetic integrity of Arctic charr (Salvelinus) populations from the Alpine region

There is a long tradition of artificially stocking lakes and rivers with fish in the hope to increase the quality and amount of fish that can be harvested. The animals used for stocking often originate in distant hatcheries or hatcheries that breed fish from remote regions. This stocking practice could have effects on the genetic integrity of resident populations. We have analysed here a case of the influence of stocking on Alpine populations of the Arctic charr (usually included into Salvelinus alpinus, but revised to Salvelinus umbla Kottelat 1997 ) within a unique geographical and historical setting. The Königssee in the Bavarian Alps (Berchtesgaden) was heavily stocked several times during the last century. However, a sample of the ancestral Königssee population still exists in the Grünsee, which lies close to Königssee, but 1000 m higher. To trace the influence of stocking in Königssee we have analysed more than 300 individuals from 10 lake populations, including the source populations of the fish used for stocking. From these we have sequenced a part of the mitochodrial control region and have typed them at six microsatellite loci. The differential distribution of haplotypes, as well as assignment tests, show that the influence of stocking on the genetic integrity of the Königssee population has been negligible. However, our data reveal that in another lake included in our study (Starnberger See), the ancestral population was apparently replaced completely by the populations used for stocking. The major difference between the lakes is the relative preservation of ecological integrity. Königssee was ecologically stable in the past, whereas Starnberger See was heavily polluted at one point, with charr approaching extinction. Interestingly, in a lake neighbouring Starnberger See, the Ammersee, which was also subject to strong pollution but not stocked, the ancestral population has recovered. Our data suggest that the practice of artificial stocking should be reconsidered, or at least monitored for effectiveness.     

Growth and contribution of stocked channel catfish,Ictalurus punctatus (Rafinesque, 1818): the importance of measuring post‐stocking performance

In this study it was sought to quantify post‐stocking growth, survival, and contribution of advanced size (178 mm total length [TL]) channel catfish Ictalurus punctatus fingerlings, something rarely done. Channel catfish populations were evaluated before (May 2010) and after (May to August 2011 and 2012) stocking. Relative abundance, stocking contribution, and growth were different (P < 0.05) in the two study impoundments (lakes Lone Chimney and Greenleaf, Oklahoma). For fish stocked in Lake Lone Chimney, stocking contribution was lower (3–35%), and average length and weight of stocked fish by age‐2 reached 230 mm TL and 85 g, whereas the stocking contribution (84–98%) and growth in length (340 mm TL) and weight (280 g) were higher by age‐2 in Lake Greenleaf. Given these unambiguous differences of post‐stocking performance, benchmark metrics that represent population‐level information such as relative abundance and average length and weight of the sample masked these significant differences, highlighting the importance of marking hatchery‐fish and then following them through time to determine the effectiveness of stocking. These results suggest that stock enhancement programmes would benefit from studies that quantify post‐stocking performance of hatchery fish.     

Growth differences between naturally recruited and stocked European eel Anguilla anguilla from different habitats in Lithuania

European eels Anguilla anguilla from freshwater lakes in Lithuania had slower growth rates and lower backcalculated total lengths ( L _T) than those from lagoons and coastal waters, but no significant differences were found among fish with different migratory histories or between naturally recruited and stocked fish except a higher L _T at age of stocked European eels at ages 5 to 8 years. The asymptotic L _T did not differ among habitats or migratory histories, but the stocked eels in the lakes had smaller K (coefficient from the von Bertalanffy growth function) than did the both naturally recruited and stocked eels in the lagoon and coastal waters. The growth rate of European eels in Lithuania might be influenced mainly by different habitats rather than different migratory histories and stocking. The lower L _T at age of naturally recruited fish at ages 5–8 years compared to stocked fish might result from the extra energy costs entailed in migration from the Atlantic and across the Baltic Sea.     

Evidence of two contrasting brown trout Salmo trutta populations spatially separated in the River Borne (France) and shift in management towards conservation of the native lineage

A multidisciplinary study was made of brown trout Salmo trutta in the Borne River, a typical fast-flowing mountain stream in the Northern French Alps, in the geographical range of the Mediterranean lineages (ML). Information on (1) the proportion of stocked fluoro-marked fish in the angling harvest, (2) the introgression of introduced DNA microsatellite alleles into the native gene pool and (3) the demography of the population in situ in autumn revealed two contrasting populations separated by a physical barrier to upstream migration. A native S. trutta population ( c . 10 000 adults) lives downstream of the barrier and is characterized by a large frequency of ML alleles (82–97%) and high densities (43–55 fish 100 m⁻²). This population is maintained predominantly by natural recruitment of juveniles (51–82%). In contrast, the upstream population is characterized by a large frequency of Atlantic lineage (AL) alleles (78–100%) and low densities (1–2 fish 100 m⁻²) and appears to be maintained by restocking (90–100%). The origins of these sharply contrasting populations appear to reflect isolation by an impassable barrier, catastrophic flooding, a downstream gradient in water quality, stocking and fishing pressure. The native downstream population has been resilient to large sudden floods and to intensive stockings of domesticated AL fish. The results of this study justify a shift in management towards conservation and rehabilitation of the native population.     

Dispersal and survival of stocked cyprinids in a small English river: comparison with wild fishes using a multi-method approach

Mark–recapture and fixed-station passive integrated transponder (PIT) telemetry were used to compare movements, distribution and survival of stocked juvenile chub Leuciscus cephalus and roach Rutilus rutilus with those of wild conspecifics. Daily activity of wild fish activity was affected by a combination of river flow and temperature, whereas stocked fishes were not influenced by environmental factors. PIT telemetry recorded exploratory movements of stocked L. cephalus immediately after stocking, a substantial number of stocked fish moved both downstream and upstream during periods of elevated flow, and proportionally more stocked fish moved during the first 6 weeks after release than later on. Proportionally more stocked fish than wild fish moved through PIT antennae, stocked L. cephalus moved greater distances than wild L. cephalus and were more widely distributed than wild fish. Minimum estimates of survival after 5 months were 50·5% for stocked R. rutilus and 28·0% for stocked L. cephalus . Ultimately, stocked cyprinids appeared to be able to cope with elevated flows and most remained in the river section local to the stocking location.     

Alteration of Nutrient Cycles and Algal Production Resulting from Fish Introductions intoMountain Lakes

The introduction of salmonid fishes into naturally fishless lakes represents one of the most prevalent environmental modifications of aquatic ecosystems in western North America. Introduced fish may alter lake nutrient cycles and primary production, but the magnitude and variation of these effects have not been fully explored. We used bioenergetics modeling to estimate the contributions of stocked trout to phosphorus (P) cycles across a wide range of fish densities in lakes of the Sierra Nevada, California. We also assessed the larger effects of fish-induced changes in phosphorus cycling on primary production using paleolimnological analyses from lakes in the southern Canadian Rockies. Our analyses showed that total P recycling by fish was independent of fish density but positively related to fish biomass in the Sierra Nevada. In lakes with fish populations maintained by continued stocking, fish recycled P at over twice the rate of those in lakes where introduced fish populations are maintained by natural reproduction and stocking has been discontinued. We estimate that P regeneration by introduced fishes is approximately equivalent to atmospheric P deposition to these lakes. Paleolimnological analyses indicated that algal production increased substantially following trout introductions to Rocky Mountain lakes and was maintained for the duration of fish presence. The results of our modeling and paleolimnological analyses indicate that introduced trout fundamentally alter nutrient cycles and stimulate primary production by accessing benthic P sources that are not normally available to pelagic communities in oligotrophic mountain lakes. These effects pose a difficult challenge for managers charged with balancing the demand for recreational fisheries with the need to maintain natural ecosystem processes. Received 28 March 2000; accepted 4 January 2001.     

The Chironomidae of the exposed zone of Øvre Heimdalsvatn

A total of 67 species of chironomids were caught in emergence traps placed at the shoreline. Additional larval records increase the number of species to be at least 75, a high number of species possibly explained by the exposed nature of the lake, which also provides habitats for lotic forms.
The species composition and relative abundance of the four main groups of Chironomidac indicate that Øvre Heimdalsvatn is a subarctic lake, with a strong oligotrophic character. The emergence period starts in mid-June and lasts to mid-September. Diversity studies show similarity with mountain lakes in France and with the Bodensee, but the equilability in Øvre Heimdalsvatn is lower.
The chironomid faunal composition of Øvre Heimdalsvatn seems to be determined by abiotic ecological factors and not the history of distribution.     

Response of benthic macroinvertebrates to whole-lake, non-native fish treatments in mid-elevation lakes of the Trinity Alps, California

Introduced fish reduce the abundance and diversity of native aquatic fauna, but the effect can be reduced in complex habitats. We manipulated fish populations in forested mountain lakes to determine whether or not fish affected benthic macroinvertebrate composition across lakes with differing habitat complexity. We compared abundance, biomass, body-length, and community structure of benthic macroinvertebrates from 16 lakes with three treatments (fish stocked, suspended stocking, fish removed) and unstocked fishless “controls”. Over 4 years, we assessed the relative importance of fish and environmental variables influencing the composition of benthic macroinvertebrates. Control lakes had the greatest overall abundance of macroinvertebrates when chironomid midges were excluded. Abundances of insects in the clinger/swimmer functional group and caddisflies were greatest in the control lakes but were primarily influenced by habitat variables including the availability of aquatic vegetation and wood. Total biomass and mean body length of macroinvertebrates were not affected by treatment. Taxon richness of macroinvertebrates was about 40% greater in the control lakes compared to the treatment lakes but did not differ among treatments. Our results suggest that fish reduce susceptible macroinvertebrate richness and abundances, but that changes associated with alterations of fish composition are confounded by other factors in complex lake habitats.     

Post‐stocking pallid sturgeon Scaphirhynchus albus growth,dispersal, and survival in the lower Missouri River

Hatchery augmentation has been used to mitigate declines in fish populations worldwide, especially for sturgeon species. Information regarding stocking success including survival, dispersal, and growth of sturgeon post‐stocking may refine sturgeon augmentation programs and facilitate recovery. Pallid sturgeon Scaphirhynchus albus populations have been supplemented by hatchery‐reared stocks for 25 years in the Missouri River, USA. We assessed survival, dispersal patterns, and growth characteristics of post‐stocked pallid sturgeon in the lower Missouri River. Hatchery‐reared pallid sturgeon stocked at age‐1 (4.1%) and > age‐1 (2.9%) were recaptured at a higher frequency than fish stocked at age‐0 (0.3%). Post‐stocking dispersal patterns suggested dispersal range increase as age increased, but individuals tended to remain in the same river segment as their stocking location. Growth rates varied by year class with younger year classes having truncated growth trajectories compared to older year classes. Post‐stock survival of pallid sturgeon varied by age‐at‐stocking and suggest age‐1 survival has declined through time. Augmentation of pallid sturgeon may benefit from considering dispersal from stocking location and by stocking older individuals which appear to have increased survival post‐stocking. A better understanding regarding environmental drivers of growth and specific habitat features used is needed to better predict optimal timing and location of future stockings.     

Hierarchical population structure and genetic diversity of lake trout (Salvelinus namaycush) in a dendritic system in Northern Labrador

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Stocking hatchery‐reared brown trout in different densities into a wild population – a comparison of growth and movement

In spring 2001 and 2002 a small stream was stocked with tagged hatchery‐reared yearling brown trout ( Salmo trutta ), in order to study their influence on the resident brown trout population. The stream was separated into six sections: two sections without stocking, two sections where stocking doubled the trout population and two sections where the fish population was quadrupled. The working hypothesis was that due to food limitation (competition) growth of the wild fish will be negatively influenced by stocking, and wild fish will be displaced by the (possibly more aggressive) hatchery fish. Surprisingly, growth rate of wild and stocked fish of the same age was similar and independent of stocking density. Two main reasons may be responsible for this finding: only a low percentage of the stocked fish remained in the stream, and food was not limited during summer. Only 12–19% of the stocked fish were recaptured after six months, in contrats to 40–70% of one‐year old and up to 100% of older wild trout. The wild fish were not displaced by hatchery‐reared fish: During summer the wild fish remained more or less stationary, whereas most of the stocked trout had left their release site. The results indicate that in a natural stream stocking of hatchery reared brown trout does not influence negatively growth and movement of the wild fish independent of stocking density.     

Examination of reproductive success of transplanted stocks in an amphidromous fish, Plecoglossus altivelis (Temmink et Schlegel) using mitochondrial DNA and isozyme markers.

Isozymes and mitochondrial DNA (mtDNA) were used as genetic markers to evaluate the reproductive success of transplanted stocks of ayu Plecoglossus ultivelis in a highly stocked Japanese river, the Shinano. Two major alleles at the MPP and GPI-A * loci and 26 mtDNA clonal genotypes were demonstrated and surveyed in two non-native source populations and five collections from the Shinano River. Contributing populations were clearly discriminated in their isozymes and mtDNA composition. Despite the stocking of large numbers of non-native fish, they have had little effect on the genetic characteristics of the native population. Maturation and migration of the stocking and native populations was traced by genetic markers to clear the mechanism of this event. All the contributing populations join together in the migration toward the spawning ground throughout the spawning season but these contributing populations occur independently of each other in other sites of the river.     

Whitefish stocking in acidified lakes: ecological and physiological responses

In autumn 1986, six small lakes at different stages of acidification were stocked with one-summer-old whitefish, Coregonus pallasi Valenciennes 1848, in order to see whether whitefish stocking would be a suitable method for the mitigation of acidification effects. In two of the lakes the introduction was a complete failure: the whitefish did not survive, evidently due to high acidity and high aluminium concentrations of the lake waters. In one of the most acidified lakes (pH 4.3–4.8, Al_lab 29–125 g 1^–1) and in two less acidic lakes (pH 5.0–5.2 and 5.4–6.4), introduction was successful. Three years after the introduction, the mean weights of the fish in those three lakes were 580, 250 and 360 g respectively, with the weight and also the condition factor of stocked whitefish being highest in the most acidified lake. In that lake there were few or no fish present during the introduction, whereas in the less acid lakes there were dense populations of perch and therefore a potential interspecific competition for food. Different availability of food in the lakes was presumed to be the main reason for the growth differences. Plasma Na⁺ and Cl^– concentrations of whitefish were lower in the acidic lakes than in the lake with pH around 6 three years after stocking. This suggests that, despite the good growth and highest condition factor of whitefish in the most acid lake, the fish still experienced some acid stress.     

The effect of stocking with 0+ year age‐class Atlantic salmon Salmo salar fry: a case study from the River Bush,Northern Ireland

An enhancement programme based on stocking 0+ year age‐class Atlantic salmon Salmo salar, conducted in the River Bush, Northern Ireland, U.K. over the period 1996–2005, was reviewed with reference to the performance and biological characteristics of wild fish. Wild ova to 0+ year fry (summer) survival was c. 8% with subsequent wild 0+ year fry‐to‐smolt survival c. 9%. Stocked unfed 0+ year juveniles gave c. 1% survival to smolt whilst fed 0+ year S. salar stocked in late summer exhibited survival at c. 5%. Stocking with unfed and fed fry contributed to increased smolt production and helped attain local management objectives between 2001 and 2005. Significant differences in biological characteristics were observed between wild and stocked‐origin fish. Wild‐smolt cohorts were dominated by 2+ year age‐class fish on the River Bush whilst smolts originating from fed fry mostly comprised younger 1+ year individuals. The mean mass of 1+ year smolts derived from stocked fed fry was significantly lower than that of wild 1+ year smolts, although these differences were not evident between older age classes. Differences in run timing between wild smolts and smolts derived from stocked fry were also apparent with the stocked‐origin fish tending to run earlier than wild fish. Although the stocking exercise was useful in terms of maximizing freshwater production, concerns over the quality of stocked‐origin recruits and the long term consequences for productivity are highlighted.     

Population collapses in introduced non-indigenous crayfish

Invasive species often have instable population dynamics and are known to collapse or oscillate heavily after passing through the initial lag/growth phases. Long-term data-series documenting these fluctuations are however rare. We use long-term (starting in the early 1960s), semi-quantitative data on the invasive signal crayfish (Pacifastacus leniusculus), capturing its population development after introduction in 44 Swedish lakes. In total 18 (41 %) of these populations had experienced a collapse. A stepwise discriminant function analysis including 20 different ecological or physicochemical characteristics identified three variables explaining collapses in the following order: stocking year, population age and mean air temperature. Populations stocked in the 1980s were more likely to collapse than populations stocked in the 1970s. Lakes with collapses were located in areas with 0.4 °C higher yearly mean air temperatures than the still viable populations. Collapses also depended on the time phase of the population and started to occur 12 years after stocking and were most frequent in the interval 16–20 years after stocking and after 11–15 years duration of the established phase with harvestable densities. An analysis of prevalence and pathogen load of Aphanomyces astaci was conducted in eight of the studied populations. A. astaci was present in all populations but neither the level of prevalence nor the pathogen load in infested specimens differed significantly between lakes with collapses and lakes without. Our results highlight the potential sensitivity and instability of introduced crayfish. The importance of density-dependence and temperature suggest that both climate variability and/or fisheries can influence these processes.     

An examination of the post-planting success of hatchery brook charr possessing different mitochondrial DNA haplotypes

Previous mtDNA and allozyme studies of brook charr Salvelinus fontinalis from northern Ontario lakes that had been extensively stocked with a government hatchery strain, yielded results that were incongruous with a neutral theory of selection in the mtDNA genome. A single haplotype (haplotype 1) predominated in all the fish sampled from these lakes, which is consistent with the observed distribution of this haplotype in northern portions of the species range. However, haplotype 1 does not occur in a high frequency within the hatchery strain. In a direct survivorship experiment, of 1500 fish stocked into an Ontario lake in the spring of 1994 containing a native population of brook charr, no significant differences between stocking and recapture mtDNA distributions were observed in the summer of 1994. However, a survey of young of the year in the following spring did not reveal the presence of any 'private' hatchery haplotypes. Significant differences in the distribution of mtDNA haplotypes were observed in two stocked lakes compared to those in the putative source hatchery strain. Haplotype 1 predominated in both these stocked lakes, suggesting that fish with this haplotype possessed increased survivorship.     

Stocking impact and temporal stability of genetic composition in a brackish northern pike population (Esox lucius L.), assessed using microsatellite DNA analysis of historical and contemporary samples

During the last decade, brackish northern pike populations in Denmark have been subject to stocking programmes, using nonindigenous pike from freshwater lakes, in order to compensate for drastic population declines. The present study was designed to investigate the genetic impact of stocking freshwater pike into a brackish pike population in Stege Nor, Denmark. We analysed polymorphism at eight microsatellite loci in samples representing the indigenous Stege Nor population prior to stocking (ie from 1956 to 1957), along with a sample of the contemporary Stege Nor population and samples from the three populations used for stocking. Despite large numbers of stocked fry, the results from both individual and population level admixture analyses demonstrated extremely poor performance and <1% introgression of stocked freshwater pike into the brackish pike population. Furthermore, pairwise F(ST) estimates between samples demonstrated close genetic relationship among temporal samples from Stege Nor, indicating temporal stability over the last 45 years. We also estimated the effective population size (N(e)) of pike in Stege Nor and applied a test for recent population bottlenecks. The harmonic mean of N(e) was relatively high (>250), but there were indications of bottlenecks in all samples and populations. We ascribe this finding to historical rather than recent bottlenecks, possibly dating back to founder events associated with postglacial recolonisation.