Conservation of endemic landlocked salmonids in regulated rivers: a case‐study from Lake Vänern,Sweden

Conservation of migratory salmonids requires understanding their ecology at multiple scales, combined with assessing anthropogenic impacts. We present a case‐study from over 100 years of data for the endemic landlocked Atlantic salmon (Salmo salar, Salmonidae) and brown trout (Salmo trutta, Salmonidae) in Lake Vänern, Sweden. We use this case‐study to develop life history‐based research and monitoring priorities for migratory salmonids. In Vänern, small wild populations of salmon and trout remain only in the heavily regulated Rivers Klar (Klarälven) and Gullspång (Gullspångsälven), and commercial and sport fisheries are maintained by hatchery stocking. These populations represent some of the last remaining large‐bodied (up to 20 kg) landlocked salmon stocks worldwide. We found that one of four stocks of wild fish has increased since 1996; the other three remain critically low. Hatchery return rates for three of four stocks appear stable at roughly 1% and annual fisheries catch is roughly 75 metric tons, with an estimated 7.5% of hatchery smolts being recruited to the fishery; this also appears relatively stable since 1990. Our analysis reveals much uncertainty in key data requirements, including both river return and fisheries catch rates, estimates of wild smolt production and survival, and hatchery breeding and genetics protocols. These uncertainties, coupled with a lack of information on their riverine and lacustrine ecology, preclude effective management of these unique populations. We conclude with a framework for a life history‐based approach to research and monitoring for Vänern salmon and trout, which should be applicable for all endemic, migratory salmonid populations.     

The response of a brown trout stocks and perception of anglers to cessation of brown trout stocking

Abstract Since the mid‐1980s, water quality has improved in German rivers, and fisheries have recovered. However, it is unsure whether the improvements are sufficient to maintain stocks and angling yield without supplementary stocking. This study examined the development in the local brown trout, Salmo trutta L., stocks and angling yield in the River Wutach following cessation of stocking in 2001, using electric fishing and an angler questionnaire. Natural reproduction was recorded in each year between 2001 and 2007, and a stable stock of trout >20 cm was found. Trout catches by anglers increased after stocking ceased and approximately 60% of the anglers were convinced that stocking was unnecessary to maintain the brown trout stocks or angler catch satisfaction.     

Are growth and recapture of hatchery‐reared and resident brown trout (Salmo trutta L.) density dependent after stocking?

Abstract –  The population density of brown trout ( Salmo trutta ) in a small natural system was manipulated in six equal-length stream sections by stocking hatchery-reared 1+ brown trout (unstocked, tripled and quintupled) over two consecutive years. The results showed that hatchery-reared trout grew more slowly and were more mobile than resident trout, and that their growth was inversely density dependent. In contrast, growth of the resident trout was density independent. The recapture of 1+ resident and hatchery-reared trout was inversely density dependent. This is most likely a consequence of increased competition. However, after a single winter the population density returned to its base level prior stocking and older resident trout showed no density-dependent recapture. Thus, the advantage of stocking, here higher biomass, may have the detrimental effect of decreasing resident stocks of the same size class during summer.     

Effects of stocking on the genetic structure of brown trout,Salmo trutta,in Central Europe inferred from mitochondrial and nuclear DNA markers

Abstract Stocking has had a considerable effect on wild brown trout, Salmo trutta L., populations throughout Europe. To elucidate this impact and to outline further management strategies, the genetic structure of 25 wild populations and five hatchery stocks from Czech Republic and Slovakia were analysed using mitochondrial (control region) and nuclear DNA (microsatellites, LDH‐C1*) markers. Stocking practices have caused massive hybridisation between the Atlantic and Danube brown trout strains in the central Danube basin and have lead to a loss of among‐population divergence in Slovakia and the eastern part of Czech Republic. Comparison with studies from neighbouring countries revealed substantial differences in haplotype, allele frequencies and genetic diversity across Central Europe. Differences in stocking management and origin of breeding stocks appear to be crucial factors for the spatial variability of the genetic structure of brown trout.     

Effect of supplementary stocking of juvenile brown trout,Salmo trutta,on yield in a Norwegian mountain reservoir

Abstract The effect of supplementary stocking of juvenile (age 0+), hatchery‐reared, brown trout, Salmo trutta L., on annual yields was assessed in a Norwegian mountain reservoir between 1979 and 2007. Fishing was mainly carried out by local fishermen with benthic gillnets. During the study period, annual stocking ranged from 0 to 52 500 fish (19.8 ha^?1). No stocking has been carried out since 1997. Annual yield varied from 1650 to 5653 kg, corresponding to 0.62–2.13 kg ha^?1. Exploitation rate in terms of number of gillnets and mean weight of 6+ fish (age when catchable size was reached) explained 64% of the variability in catches. Stocked fish contributed very little to the yield or catch‐per‐unit‐effort, exhibiting no positive correlation with stocking density. The lack of contribution from stocked fish was probably caused by a competitive bottleneck in the eroded epibenthic zone, causing high juvenile mortality. If stocking continues, it is recommended that fish with body lengths >15–20 cm are used.     

Empirical move‐on rules to inform fishing strategies: a New England case study

Increasingly, fisheries are being managed under catch quotas that are often further allocated to specific permit holders or sectors. At the same time, serious consideration is being given to the effects of discards on the health of target and non‐target species. Some quota systems have incorporated discard reduction as an objective by counting discards (including unmarketable fish) against the overall quota. The potential effect of the introduction of a quota system that includes accountability for discards on the fishing strategies employed by fishermen is enormous. This is particularly true for multispecies fisheries where healthy and depleted stocks co‐exist; resulting in a trip's catch being applied to very large and very small stock quotas simultaneously. Under such a scenario, fishermen have a strong incentive to minimize (i) catch of low‐quota or ‘choke’ stocks, (ii) regulatory discards due to minimum size limits and (iii) catch partially consumed by predators. ‘Move‐on’ rules (i.e. event‐triggered, targeted, temporary closure of part of a fishery when a catch or bycatch threshold is reached) have been employed in a variety of fisheries. However, their efficacy has been limited by a lack of empirical analyses underpinning the rules. Here, we examine the utility of spatiotemporal autocorrelation analyses to inform ‘move‐on’ rules to assist a sector of the New England Multispecies Fishery to reduce discards and maximize profits. We find the use of empirical move‐on rules could reduce catch of juvenile and choke stocks between 27 and 33%, and depredation events between 41 and 54%.     

Use of first‐order tributaries by brown trout (Salmo trutta) as nursery habitat in a cold water stream network

Many investigators have examined the importance of suitable in‐stream habitat and flow regime to salmonid fishes. However, there is much less known about the use of small (<5 l·s^?1 discharge) first‐order streams within a larger stream network by salmonids. The purpose of this study was to evaluate the use of small headwater streams by juvenile brown trout Salmo trutta in the Emmons Creek stream network in Wisconsin, USA, and to determine whether abundance was related to habitat variables in these streams. Fishes in eight spring‐fed first‐order streams were sampled during a 7‐month period using a backpack electroshocker and measured for total length. Habitat variables assessed included stream discharge, water velocity, sediment composition and the abundance of cover items (woody debris and macrophytes). Densities of YOY trout ranged from 0 to 1 per m² over the course of the study and differed among first‐order streams. Stepwise multiple regression revealed discharge to be negatively associated with trout density in spring but not in summer. All other habitat variables were not significantly related to trout density. Our results demonstrate the viability of small first‐order streams as nursery habitat for brown trout and support the inclusion of headwater streams in conservation and stream restoration efforts.     

Controls for trout and char migratory/resident behaviour mainly in stream systems above and below waterfalls/barriers: a multidecadal and broad geographical review

Northcote TG. Controls for trout and char migratory/resident behaviour mainly in stream systems above and below waterfalls/barriers: a multidecadal and broad geographical review. Ecology of Freshwater Fish 2010: 19: 487–509. © 2010 John Wiley & Sons A/S Abstract – Controls for trout and char migratory and resident behaviour in rivers and streams for above and below natural and man‐made waterfalls/barriers are covered in a multi‐decadal (1950–2000) and wide geographical review (North America, UK, Europe, Japan) that includes reference to over 380 publications. These note 53 for rainbow trout, 61 for cutthroat trout, 104 for brown trout, 47 for bull trout, 41 for brook trout, 35 for Dolly Varden and 42 for white‐spotted char, plus a few general contributions of relevance on some. For each of these species, there has been a major increase in relevant decadal publications since the early 1980s, no doubt in large part because of the upsurge in micro‐genetic methodology for DNA and related technology, coupled with a broadening of interest in stream migratory behaviour of salmonids. Main mechanisms for the control of stream migratory versus resident behaviour appear to cover an interplay among both genetic and environmental factors; in some populations and locations, genetic controls seem to be more important than environmental ones, but in others the reverse. Habitat degradation by various human activities and their introductions of non‐native fish species are becoming causes of reduction in abundance of above and below waterfall stream populations for several of these salmonid species.     

Typology of landowners in Norwegian salmon angling: attitudes towards river owner organisations and management actions

Abstract Atlantic salmon, Salmo salar L., recreational fisheries in Norway are facing shorter seasons and harvest restrictions because of low adult migration runs. Private fishing‐right holders (landowners) are important stakeholders, being co‐managers of the stocks, owners of salmon habitat and suppliers of angling. Landowners saw measures addressing salmon farming and Gyrodactylus salaris (Malmberg) as the most important management actions to strengthen stocks and downplayed actions restricting their own activity or gain. The results showed a need to build knowledge and improve communication between landowners and river owner organisations about the effects of stocking, catch and release, and other management actions. Four distinct landowner types were identified, based on their objectives for the fishing right. The diversity in the landowner group suggests managing angling tourism will be challenging, and cooperation is required to manage salmon stocks. Policy instruments to facilitate cooperation are discussed for each landowner type.     

Genetic refuges for a self‐sustained fishery: experience in wild brown trout populations in the eastern Pyrenees

Abstract – Management policies balancing harvest and conservation of natural populations of fish are difficult to establish, both scientifically and politically. This issue is particularly difficult when those populations represent native genetic resources. Since 1997, several brown trout populations in the eastern Pyrenees Mountains (Spain) were designated as ‘genetic refuges’ under varying fishing regulations, where releases of hatchery‐origin fish are not permitted. We analysed genetic variation in samples of brown trout from six of those refuge populations and four non‐refuge populations within the same region. Each population was sampled in four separate years: 1993, 1999, 2004 and 2006. Our analyses were based on a diagnostic allele (LDH‐C*90) that distinguishes native and exogenous hatchery populations. Comparisons were based on stocking histories before and after refuge designations and on three management strategies: fished, unfished and catch‐and‐release. Overall, we detected significant genetic introgression resulting from past stocking practices despite the current restriction of hatchery releases imposed by the recent genetic refuge policy. However, this new policy has prevented detectable introgression from increasing throughout the region and together with additional measures on length and number of captured fish is contributing to self‐sustained fisheries that are achieving conservation goals. Quick acceptance of ‘genetic refuges’ by anglers in one particular river, the Ter River basin, has been a key factor in protecting native gene pools compared with the Segre River basins where refuges were not readily accepted.     

Inland fish stock assessment: Applying data‐poor methods from marine systems

Inland fisheries can be diverse, local and highly seasonal. This complexity creates challenges for monitoring, and consequently, many inland fish stocks have few data and cannot be assessed using methods typically applied to industrial marine fisheries. In such situations, there may be a role for methods recently developed for assessment of data‐poor fish stocks. Herein, three established data‐poor assessment tools from marine systems are demonstrated to highlight their value to inland fisheries management. A case study application uses archived length, catch and catch‐per‐unit‐effort data to characterise the ecological status of an important recreational brown trout stock in an Irish lake. This case study is of specific use to management of freshwater sport fisheries, but the broader purpose of the paper was to provide a crossover between marine and inland fisheries science, and to highlight accessible data‐poor assessment approaches that may be applicable in diverse inland systems.     

Non‐indigenous fish in protected spaces: Trends in species distribution mediated by illegal stocking

1. Many freshwater non‐indigenous species (NISs) are stocked for recreational fishing, in some cases illegally in protected areas. In this study, fish communities were monitored using environmental DNA, electrofishing and anglers’ catches as the sources of samples in a mountainous Biosphere Reserve in Asturias (northern Spain), where stocking is forbidden.
2. Three NISs have been introduced illegally in the protected area and have shown increasing populations in the last two decades. Two species used as fishing bait, Squalius carolitertii (chub) and Phoxinus phoxinus (minnow), are expanding in running waters. Oncorhynchus mykiss (rainbow trout) was also detected and is likely to have been introduced for angling or from fish farm escapes.
3. The results suggest that sustained illegal stocking contributed to the increase of the three NISs. In contrast, Salmo trutta (brown trout) of northern European lineages, identified from *90 alleles at the LDH‐C1 locus, and formerly legally stocked for angling, is decreasing, most likely as a result of climate change. Climate change could also contribute to the expansion of the two non‐indigenous cyprinids to colder upstream areas.
4. Through the application of a social survey, it was found that unlike other population groups, anglers in the region significantly preferred stocking over environmental improvement for the management of fish populations. The results obtained suggest that raising the awareness of anglers about the importance of safeguarding native fish species could help to prevent the spread of NISs in protected areas.

     

Genetic persistence of an initially introduced brown trout (Salmo trutta L.) population despite restocking of foreign conspecifics

The persistence of the initial genetic structure despite later stocking with foreign conspecifics is influenced by several factors, leading to different levels of introgression. Stream spawning has been assumed to be the prevailing recruitment strategy of brown trout (Salmo trutta L.). However, in lakes with limited stream spawning habitat, but still with high natural recruitment, lake spawning has been proposed. Using fourteen microsatellites, we assessed the genetic structure in a small allopatric brown trout population in the Lake Skavatn, located on the Hardangervidda mountain plateau, Norway. A total of 265 brown trout were obtained from the lake itself, the outlet stream, two tributaries, a littoral sample from a presumed lake spawning site, a suggested stocking source, and a lake sample from 1967, representing the initial lake population. The six Skavatn samples were best represented by three genetic components, showing a shift from the initial population, but with no genetic signal from the suggested stocking source. The littoral sample had the largest similarity to the initial lake population, possibly indicating the importance of lake spawning in a system where streams offer unpredictable spawning and rearing conditions. Due to large annual variations in recruitment contributions from the different spawning and rearing locations, the genetic structure of the lake population probably vary over time.     

The balancing act of captive breeding programmes: salmon stocking and angler catch statistics

The debate over Atlantic salmon, Salmo salar L., stocking in Britain centres on the trade‐off between enhancing rod fisheries and harming wild populations. This article informs the debate by quantifying the relationship between stocking and angler catch statistics for 62 rivers over 15 years. After controlling for environmental factors affecting adult abundance, the 42 rivers with stocking had non‐significantly lower mean catch statistics than the 20 rivers without stocking. This difference increased with the age of stocked fish. Among stocked rivers, weak relationships between mean stocking effort and catch statistics also became more negative with the age of stocked fish. For stocked rivers, there was no evidence for a generally positive relationship between annual stocking efforts and catch statistics. Those rivers for which stocking appeared to improve annual rod catches tended to have lower than expected mean rod catches. The results suggest the damage inflicted on wild salmon populations by stocking is not balanced by detectable benefits to rod fisheries.     

Effect of fine wood on juvenile brown trout behaviour in experimental stream channels

In‐stream wood can increase shelter availability and prey abundance for stream‐living fish such as brown trout, Salmo trutta, but the input of wood to streams has decreased in recent years due to harvesting of riparian vegetation. During the last decades, fine wood (FW) has been increasingly used for biofuel, and the input of FW to streams may therefore decrease. Although effects of in‐stream FW have not been studied as extensively as those of large wood (LW), it is probably important as shelter for small‐sized trout. In a laboratory stream experiment, we tested the behavioural response of young‐of‐the‐year wild brown trout to three densities of FW, with trout tested alone and in groups of four. Video recordings were used to measure the proportion of time allocated to sheltering, cruising and foraging, as well as the number of aggressive interactions and prey attacks. Cruising activity increased with decreasing FW density and was higher in the four‐fish groups than when fish were alone. Foraging decreased and time spent sheltering in FW increased with increasing FW density. Our study shows that juvenile trout activity is higher in higher fish densities and that trout response to FW is related to FW density and differs from the response to LW as reported by others.     

Analysis of Fishery Records from Cow Green Reservoir, Upper Teesdale, 1971–1980

Apart from some irregularities reflecting changes in permit costs (1975) and the effects of drought (1976), there were increases in angler-visits, catch per angler-visit and number of fish caught during the first 5 years of the fishery. Thereafter, there were about 1000–1500 angler-visits per season, a mean catch of 1.0–1.2 trout per angler-visit and an annual catch of 1200–1800 trout. Between 1971 and 1980 the mean weight of the fish caught fell from 472 to 349 g. During the period 1976–1980 the mean catch per angler-visit was 1.2 fish. This compares favourably with results from a selection of upland and lowland British reservoirs. The mean weight of individual trout caught was 362 kg and this value is typical of both stocked and unstocked upland reservoirs. The mean annual catch during 1976–1980 was about 2 kg ha^?1. This is similar to the values observed in other unstocked upland reservoirs. The gross yield from such reservoirs can be increased by stocking but the limited data available suggest that at stocking rates above about 5 kg ha^?1 the net yield (i.e. weight caught — weight stocked) becomes negative.     

Electrophoretic evaluation of a supplemental stocking programme for brown trout, Salmo trutta L.

Abstract. Over a 15-year period hatchery brown trout, Salmo trutta L., have been added 10 Lower Lough Erne, Northern Ireland to supplement declining native populations. Introductions have mainly comprised eyed ova and fingerlings, stocked into a number of rivers in the Erne drainage. Utilizing a natural genetic tag an electrophoretic assessment of the stocking programme was undertaken. The percentage hatchery genetic contribution in trout populations varied widely from river to river (19%-91%). Lough-caught brown trout (3+ and older) showed a substantial (21·5%) hatchery genetic component. Introgression of native-and hatchery stocks was evident. The resultant deleterious genetic consequences for the conservation of the unique Lough Erne brown trout gene pool arc discussed and alternative management strategies are proposed.     

Competitive interactions among multiple non‐native salmonids and two populations of Atlantic salmon

Competitive interactions with non‐native species can have negative impacts on the conservation of native species, resulting in chronic stress and reduced survival. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (Sebago and LaHave) that are being used for reintroduction into Lake Ontario were placed into semi‐natural stream tanks with four non‐native salmonid competitors that are established in Ontario streams: brown trout (S. trutta), rainbow trout (Oncorhynchus mykiss), Chinook salmon (O. tshawytscha) and coho salmon (O. kisutch). Brown trout and rainbow trout reduced the survival and fitness‐related traits of Atlantic salmon, whereas Chinook salmon and coho salmon had no impact on these traits. These data support theories on ecological niche overlap and link differences in observed aggression levels with competitive outcomes. Measurements of circulating hormones indicated that the Atlantic salmon were not chronically stressed nor had a change in social status at the 10‐month time point in the semi‐natural stream tanks. Additionally, the Sebago population was better able to coexist with the non‐native salmonids than the LaHave population. Certain populations of Atlantic salmon may thus be more suitable for some environments of the juvenile stream phase for the reintroduction into Lake Ontario.     

Benefits of repeated stocking with adult,hatchery‐reared brown trout,Salmo trutta,to recreational fisheries?

Abstract  Recaptures of adult, hatchery-reared, brown trout, Salmo trutta L., and fishing time from anglers were used to evaluate the benefits of stocking programmes with repeated releases of adult brown trout. The recapture rate varied between 17% and 29%. The time between stocking and capture (referred to as residence time) varied between 1 and 160 days (median 3–49 days). Between 67% and 84% of trout caught in the river were recently released fish. Fishing effort increased after stocking, thereby increasing the impact of angling on wild stocks. Stocking with adult brown trout decreases the impact of angling on wild trout only if the time spent fishing by all anglers is kept stable. Furthermore, because of the short residence time of stocked trout, long-term impacts through competition for space and food, or genetic impact through introgression, are limited.     

Can catch share fisheries better track management targets?

Fisheries management based on catch shares – divisions of annual fleet‐wide quotas among individuals or groups – has been strongly supported for their economic benefits, but biological consequences have not been rigorously quantified. We used a global meta‐analysis of 345 stocks to assess whether fisheries under catch shares were more likely to track management targets set for sustainable harvest than fisheries managed only by fleet‐wide quota caps or effort controls. We examined three ratios: catch‐to‐quota, current exploitation rate to target exploitation rate and current biomass to target biomass. For each, we calculated the mean response, variation around the target and the frequency of undesirable outcomes with respect to these targets. Regional effects were stronger than any other explanatory variable we examined. After accounting for region, we found the effects of catch shares primarily on catch‐to‐quota ratios: these ratios were less variable over time than in other fisheries. Over‐exploitation occurred in only 9% of stocks under catch shares compared to 13% of stocks under fleet‐wide quota caps. Additionally, over‐exploitation occurred in 41% of stocks under effort controls, suggesting a substantial benefit of quota caps alone. In contrast, there was no evidence for a response in the biomass of exploited populations because of either fleet‐wide quota caps or individual catch shares. Thus, for many fisheries, management controls improve under catch shares in terms of reduced variation in catch around quota targets, but ecological benefits in terms of increased biomass may not be realized by catch shares alone.