The anadromous life cycle of Atlantic salmon Salmo salar involves long migrations to novel environments and challenging physiological transformations when moving between salt-free and salt-rich waters. In this article, (1) environmental factors affecting the migration behaviour and survival of smolts and post-smolts during the river, estuarine and early marine phases, (2) how behavioural patterns are linked to survival and (3) how anthropogenic factors affect migration and survival are synthesized and reviewed based on published literature. The timing of the smolt migration is important in determining marine survival. The timing varies among rivers, most likely as a consequence of local adaptations, to ensure sea entry during optimal periods. Smolts and post-smolts swim actively and fast during migration, but in areas with strong currents, their own movements may be overridden by current-induced transport. Progression rates during the early marine migration vary between 0.4 and 3.0 body lengths s(-1) relative to the ground. Reported mortality is 0.3-7.0% (median 2.3) km(-1) during downriver migration, 0.6-36% (median 6.0) km(-1) in estuaries and 0.3-3.4% (median 1.4) km(-1) in coastal areas. Estuaries and river mouths are the sites of the highest mortalities, with predation being a common cause. The mortality rates varied more among studies in estuaries than in rivers and marine areas, which probably reflects the huge variation among estuaries in their characteristics. Behaviour and survival during migration may also be affected by pollution, fish farming, sea lice Lepeophtheirus salmonis, hydropower development and other anthropogenic activities that may be directly lethal, delay migration or have indirect effects by inhibiting migration. Total mortality reported during early marine migration (up to 5-230 km from the river mouths) in the studies available to date varies between 8 and 71%. Hence, the early marine migration is a life stage with high mortalities, due to both natural and human influences. Factors affecting mortality during the smolt and post-smolt stages contribute to determine the abundance of spawner returns. With many S. salar populations in decline, increased mortality at these stages may considerably contribute to limit S. salar production, and the consequences of human-induced mortality at this stage may be severe. Development of management actions to increase survival and fitness at the smolt and post-smolt stages is crucial to re-establish or conserve wild populations.
We review factors affecting the withinriver spawning migration of Atlantic salmon. With populations declining across the entire distribution range, it is important that spawners survive in the last phase of the spawning migration. Knowledge on the factors affecting migration is essential for the protection of populations, and to increase the success of reintroduction programmes. A number of studies have documented that the upstream migration may be delayed for many weeks at man-made obstacles such as power station outlets, residual flow stretches, dams, weirs and fishways. The fish may also be delayed at natural migration barriers. Often, the magnitude of delay is not predictable; fish may be considerably delayed at barriers that appear to humans to be easily passable, or they may quickly pass barriers that appear difficult. Stressful events like catch-and-release angling may affect upstream migration. Impacts of human activities may also cause altered migration patterns, affect the withinriver distribution of the spawning population, and severe barriers may result in displacement of the spawning population to other rivers. Factors documented to affect within-river migration include previous experience, water discharge, water temperature, water velocity, required jump heights, fish size, fish acclimatisation, light, water quality/pollution, time of the season, and catch and handling stress. How each of these factors affects the upstream migration is to a varying extent understood; however, the effects may differ among different river sections and sites. There are likely a number of additional important factors, and the relationship between different factors is complex. The understanding of general mechanisms stimulating fish within-river migration are still lacking, and it cannot be reliably predicted under which conditions a fish will pass a given migration barrier or which conditions are needed to stimulate migration at different sites. The strong focus on the effects of water discharge in past work may have hampered consideration of other factors. Exploration of the influence of these other factors in future studies could improve our understanding of what controls the upstream migration.
The escape of fish from aquaculture is perceived as a threat to wild fish populations. The escapes problem is largely caused by technical and operational failures of fish farming equipment. In Norway ), despite the total number of salmon held in sea-cages increasing by 44% during this period. No similar decrease in escaped cod has occurred, suggesting that other measures, such as improved netting materials for sea-cages, are required. In addition to escaping as juveniles or adults, cod may reproduce in seacages, and thus fertilised eggs escape to the environment. The ecological effects of 'escape through spawning' are unclear, and methods to inhibit escape by this mechanism are being explored. To prevent escapes of juvenile and adult fish as sea-cage aquaculture industries develop, we recommend that policy-makers implement a 5 component strategy: (1) establish mandatory reporting of all escape incidents; (2) establish a mechanism to analyse and learn from the mandatory reporting; (3) conduct mandatory, rapid, technical assessments to determine the causes of escape incidents involving more than 10 000 fish; (4) introduce a technical standard for sea-cage aquaculture equipment coupled with an independent mechanism to enforce the standard; and (5) conduct mandatory training of fish farm staff in escape-critical operations and techniques.
Considerable technical developments over the past half century have enabled widespread application of electronic tags to the study of animals in the wild, including in freshwater environments. We review the constraints associated with freshwater telemetry and biologging and the technical developments relevant to their use. Technical constraints for tracking animals are often influenced by the characteristics of the animals being studied and the environment they inhabit. Collectively, they influence which and how technologies can be used and their relative effectiveness. Although radio telemetry has historically been the most commonly used technology in freshwater, passive integrated transponder (PIT) technology, acoustic telemetry and biologgers are becoming more popular. Most telemetry studies have focused on fish, although an increasing number have focused on other taxa, such as turtles, crustaceans and molluscs. Key technical developments for freshwater systems include: miniaturization of tags for tracking small-size life stages and species, fixed stations and coded tags for tracking large samples of animals over long distances and large temporal scales, inexpensive PIT systems that enable mass tagging to yield population-and community-level relevant sample sizes, incorporation of sensors into electronic tags, validation of tag attachment procedures with a focus on maintaining animal welfare, incorporation of different techniques (for example, genetics, stable isotopes) and peripheral technologies (for example, geographic information systems, hydroacoustics), development of novel analytical techniques, and extensive international collaboration. Innovations are still needed in tag miniaturization, data analysis and visualization, and in tracking animals over larger spatial scales (for example, pelagic areas of lakes) and in challenging environments (for example, large dynamic floodplain systems, under ice). There seems to be a particular need for adapting various global positioning system and satellite tagging approaches to freshwater. Electronic tagging provides a mechanism to collect detailed information from imperilled animals and species that have no direct economic value. Current and future advances will continue to improve our knowledge of the natural history of aquatic animals and ecological processes in freshwater ecosystems while facilitating evidence-based resource management and conservation.
Atlantic salmon (Salmo salar) is an economically and culturally important species. Norway has more than 400 watercourses with Atlantic salmon and supports a large proportion of the world's wild Atlantic salmon. Atlantic salmon are structured into numerous genetically differentiated populations, and are therefore managed at the population level. Long-distance migrations between freshwater and ocean habitats expose Atlantic salmon to multiple threats, and a number of anthropogenic factors have contributed to the decline of Atlantic salmon during the last decades. Knowledge on the relative importance of the different anthropogenic factors is vital for prioritizing management measures. We developed a semi-quantitative 2D classification system to rank the different anthropogenic factors and used this to assess the major threats to Norwegian Atlantic salmon. Escaped farmed salmon and salmon lice from fish farms were identified as expanding population threats, with escaped farmed salmon being the largest current threat. These two factors affect populations to the extent that they may be critically endangered or lost, with a large likelihood of causing further reductions and losses in the future. The introduced parasite Gyrodactylus salaris, freshwater acidification, hydropower regulation and other habitat alterations were identified as stabilized population threats, which have contributed to populations becoming critically endangered or lost, but with a low likelihood of causing further loss. Other impacts were identified as less influential, either as stabilized or expanding factors that cause loss in terms of number of returning adults, but not to the extent that populations become threatened. Management based on population specific reference points (conservation limits) has reduced exploitation in Norway, and overexploitation was therefore no longer regarded an important impact factor. The classification system may be used as a template for ranking of anthropogenic impact factors in other countries and as a support for national and international conservation efforts.
Electromyogram (EMG) telemetry studies that involve remotely monitoring the locomotory activity and energetics of ¢sh are contributing important information to the conservation and management of ¢sheries resources. Here, we outline the development of this rapidly evolving ¢eld and formulate the studies conducted that utilize this technology. To date, more than 60 studies have been conducted using EMG telemetry that spans 18 species. Several general trends were observed in the methodology of the studies that we have highlighted as standards that should be adopted associated with transmitter customization, electrode placement and surgical technique. Although numerous studies have been methodological, there are still some de¢ciencies in our basic understanding of issues such as the need for individual calibration and the method of reporting or transforming data. Increasingly, this technology is being applied to address issues in conservation, management and aquaculture production. At present, the technology has been most frequently applied to the study of animal activity or energetics and to migration. Several recent studies have also focused on addressing more basic questions in ecological and evolutionary biology (e.g. parental care dynamics) similar to the large body of literature that has been collected for other taxa (e.g. marine mammals, birds), using activity telemetry. Collectively, studies conducted using EMG telemetry have contributed important information on free-swimming ¢sh that was previously di¤cult to obtain. EMG telemetry is particularly e¡ective for examining behaviour at temporal and spatial scales that are di¤cult using other techniques. The development of an ultrasonic transmitter based on the same proven principles as those used in the current radio transmitter technology will permit studies in other environments (i.e. marine, brackish, deep water) and on di¡erent species of ¢sh.We encourage the continued development and re¢nement of devices for monitoring the activity and energetics of free-swimming ¢sh, and also encourage researchers to consider EMG telemetry as a tool for addressing questions that are not e¡ectively answered with other techniques.
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