Direct evidence of increased natural mortality of a wild fish caused by parasite spillback from domestic conspecifics.

Parasite spillback from domestic animals can distort the balance between host and parasites in surrounding wildlife, with potential detrimental effects on wild populations. In aquatic environments, parasite spillback from aquaculture to wild salmon is one of the most contentious sustainability debates. In a 19 year time series of release group studies of Atlantic salmon, we demonstrated that (i) the effect of subjecting out-migrating salmon smolts to parasite treatment on marine survival has been reduced over a time, (ii) the relation between salmon lice levels in the out-migration route of the salmon and effect of treatment against the parasite is weak, but also (iii) the return rates in both treated and untreated groups of salmon are negatively correlated with salmon lice levels, and (iv) returns of wild salmon to the region are similarly negatively correlated with salmon lice levels during the out-migration year. Our study suggests that salmon lice can have a large effect on wild salmon populations that is not revealed with randomized control trials using antiparasitic drugs. This should be better accounted for when considering the impacts of farms on wild salmon populations.

The paradox of predation studies.

Understanding the causal relationships that contribute to mortality in populations is a priority for epidemiology, animal husbandry and ecology. Of all the sources of mortality in nature, predation is perhaps the most important, while simultaneously being one of the most difficult to study and understand. In this opinion piece, we use the epidemiological concept of the sufficient-component cause model to outline why we believe that predation studies often misrepresent predators as sufficient cause of death (or natural mortality) in ecological studies. This is pivotal in conservation biology because such studies have often led to demands for predator removal throughout the world. We use the sufficient-component cause model to illustrate the paradox that multiple studies, each studying singular putative causes of mortality (including predation), will sum to more than 100% mortality when added together. We suggest that the sufficient-component framework should be integrated into both fundamental and applied ecology to better understand the role of predators in natural ecosystems.

Effects of tag type and surgery on migration of Atlantic salmon (Salmo salar) smolts.

Tagging salmon smolts to provide information about the timing of outmigration has been a common approach to monitor phenology and model the risk of encountering stressors. However, the validity of tagging has come under scrutiny because of the sensitivity of this parameter in various management systems. We studied the probability of migration, timing of migration and growth during migration for Atlantic salmon smolts tagged with three different tags in the River Dale, western Norway. Two groups were tagged with passive integrated transponder (PIT) tags via a small ventral nonsurgical incision, either a 12 mm or a new 16 mm PIT tag. Two groups were subjected to surgical implantation of either a dummy acoustic transmitter or a 12 mm PIT tag (a sham surgery). Overall, 71% of the tagged smolts were recaptured at the downstream Wolf trap. Smolts from the sham tagged group were recaptured most frequently (78%) compared to dummy acoustic transmitters and 16 mm PIT tags (both 68%), but the differences were not significant. Results agree with prior assessments that longer smolts migrated earlier, with about half a day earlier migration for each millimetre total length of the smolt, but did not suggest any difference in time of migration among the tag types. Growth in length was evident from release to recapture, with smaller smolts exhibiting greater growth and no effect of tagging treatment. Our findings suggest that inferences about the timing of outmigration for salmon smolts based on acoustic tagging should be made cautiously because of the relationship among tag size, suitable fish size and the timing of a tagged individual's migration.

Salmon lice increase the age of returning Atlantic salmon.

The global increase in the production of domestic farmed fish in open net pens has created concerns about the resilience of wild populations owing to shifts in host-parasite systems in coastal ecosystems. However, little is known about the effects of increased parasite abundance on life-history traits in wild fish populations. Here, we report the results of two separate studies in which 379 779 hatchery-reared Atlantic salmon smolts were treated (or not) against salmon lice, marked and released. Adults were later recaptured, and we specifically tested whether the age distribution of the returning spawners was affected by the treatment. The estimates of parasite-induced mortality were 31.9% and 0.6% in the River Vosso and River Dale stock experiments, respectively. Age of returning salmon was on average higher in untreated [corrected] versus untreated fish. The percentages of fish returning after one winter at sea were 37.5% and 29.9% for the treated and untreated groups, respectively. We conclude that salmon lice increase the age of returning salmon, either by affecting their age at maturity or by disproportionately increasing mortality in fish that mature early.

Acoustic accelerometer transmitters and their growing relevance to aquatic science.

There has recently been great interest in the use of accelerometers onboard electronic transmitters to characterise various aspects of the ecology of wild animals. We review use cases and outline how these tools can provide opportunities for studying activity and survival, exercise physiology of wild animals, the response to stressors, energy landscapes and conservation planning tools, and the means with which to identify behaviours remotely from transmitted data. Accelerometer transmitters typically send data summaries to receivers at fixed intervals after filtering out static acceleration and calculating root-mean square error or overall dynamic body action of 2- or 3-axis acceleration values (often at 5-12.5 Hz) from dynamic acceleration onboard the tag. Despite the popularity of these transmitters among aquatic ecologists, we note that there is wide variation in the sampling frequencies and windows used among studies that will potentially affect the ability to make comparisons in the future. Accelerometer transmitters will likely become increasingly popular tools for studying finer scale details about cryptic species that are difficult to recapture and hence not suitable for studies using data loggers. We anticipate that there will continue to be opportunities to adopt methods used for analysing data from loggers to datasets generated from acceleration transmitters, to generate new knowledge about the ecology of aquatic animals.

Ecological regime shift in the Northeast Atlantic Ocean revealed from the unprecedented reduction in marine growth of Atlantic salmon.

Ecological regime shifts are abrupt changes in the structure and function of ecosystems that persist over time, but evidence of contemporary regime shifts are rare. Historical scale data from 52,384 individual wild Atlantic salmon caught in 180 rivers from 1989 to 2017 reveal that growth of Atlantic salmon across the Northeast Atlantic Ocean abruptly decreased following the year 2004. At the same time, the proportion of early maturing Atlantic salmon decreased. These changes occurred after a marked decrease in the extent of Arctic water in the Norwegian Sea, a subsequent warming of spring water temperature before Atlantic salmon entering the sea, and an approximately 50% reduction of zooplankton across large geographic areas of the Northeast Atlantic Ocean. A sudden decrease in growth was also observed among Atlantic mackerel in the Norwegian Sea. Our results point toward an ecosystem-scale regime shift in the Northeast Atlantic Ocean.

Negative impacts of the sea lice prophylactic emamectin benzoate on the survival of hatchery released salmon smolts in rivers.

Emamectin benzoate (EB) is a prophylactic pharmaceutical used to protect Atlantic salmon (Salmo salar) smolts migrating out of rivers and into the ocean against sea lice parasites. Randomized control trials comparing the marine survival of smolts treated with EB to a control group is used to calculate the fraction of marine mortality attributable to sea lice parasitism. However, it is assumed that there is no baseline difference in survival induced by the application of EB treatment. We used a combined laboratory and field study approach to investigate the potential impacts of EB treatment on behaviour and survival of hatchery-reared Atlantic salmon in western Norway. In aquaria experiments, EB-treated salmon smolts did not differ significantly in exploratory behaviour. Fish from treated groups responded similarly to simulated predator attack with spontaneous escape and elevated gill beat rate. Three rivers in the Osterfjord system of western Norway were selected for field experiments, Dale, Vosso, and Modalen. Dale River smolts were treated with intraperitoneal EB injections and had lower probability of detection in a wolf trap downstream of the release site than control smolts. Salmon smolts raised in the Vosso River hatchery were treated with EB delivered in their food and were detected on PIT antennas at the rivermouth of Vosso and Modalen at lower rates than control fish, but only when released at downstream sites. Calculation of risk ratios suggested that the bias in mortality caused by treatment with EB decreased the estimated survival of treated fish from an expected 18%to 46%, reducing the observable negative impact of sea lice on Atlantic salmon smolts in randomized control trials. The results suggest that estimates of the fraction of mortality attributable to sea lice may be underestimated due to lower baseline survival of treated fish caused by treatment and bring urgent attention towards a potential systematic underestimation of the impacts of sea lice on wild salmon.

Parasite induced mortality is context dependent in Atlantic salmon: insights from an individual-based model.

An individual-based model was parameterized to explore the impact of a crustacean ectoparasite (sea louse, Lepeophtheirus salmonis & Caligus spp.) on migrating Atlantic salmon smolt. The model explores how environmental and intrinsic factors can modulate the effect of sea lice on survival, growth and maturation of Atlantic salmon at sea. Relative to other effects, the parasite infestation pressure from fish farms and the encounter process emerge as the most important parameters. Although small variations in parasite-induced mortality may be masked by variable environmental effects, episodes of high infestation pressure from fish farms should be observable in wild populations of Atlantic salmon if laboratory studies accurately reflect the physiological effects of sea lice. Increases in temperature in the model negatively influenced fish survival by affecting the development time of the parasite at a rate that was not compensated for by the growth of the host. Discharge from rivers was parameterized to increase migration speed and influenced parasite induced mortality by decreasing time spent in areas with increased infestation pressure. Initial size and growth of the host was inversely related to the impact of the parasite because of size-dependent parasite-induced mortality in the early phase of migration. Overall, the model illustrates how environmental factors modulate effects on the host population by impacting either the parasite load or the relative effect of the parasite. The results suggest that linking population-level effects to parasite infestation pressure across climatic and environmental gradients may be challenging without correctly accounting for these effects.

Quantitative risk assessment of salmon louse-induced mortality of seaward-migrating post-smolt Atlantic salmon.

The Norwegian government recently implemented a new management system to regulate salmon farming in Norway, aiming to promote environmentally sustainable growth in the aquaculture industry. The Norwegian coast has been divided into 13 production zones and the volume of salmonid production in the zones will be regulated based on salmon lice effects on wild salmonids. Here we present a model for assessing salmon louse-induced mortality of seaward-migrating post-smolts of Atlantic salmon. The model quantifies expected salmon lice infestations and louse-induced mortality of migrating post-smolt salmon from 401 salmon rivers draining into Norwegian coastal waters. It is assumed that migrating post-smolts follow the shortest path from river outlets to the high seas, at constant progression rates. During this migration, fish are infested by salmon lice of farm origin according to an empirical infestation model. Furthermore, louse-induced mortality is estimated from the estimated louse infestations. Rivers draining into production zones on the West Coast of Norway were at the highest risk of adverse lice effects. In comparison, rivers draining into northerly production zones, along with the southernmost production zone, were at lower risk. After adjusting for standing stock biomass, estimates of louse-egg output varied by factors of up to 8 between production zones. Correlation between biomass adjusted output of louse infestation and densities of farmed salmon in the production zones suggests that a large-scale density-dependent host-parasite effect is a major driver of louse infestation rates and parasite-induced mortality. The estimates are sensitive to many of the processes in the chain of events in the model. Nevertheless, we argue that the model is suited to assess spatial and temporal risks associated with farm-origin salmon lice.

First observations of saturopeaking: Characteristics and implications.

During the monitoring of total dissolved gas (TDG) saturation in the Vetlefjordelva River in western Norway in 2014-2015, characteristic waves of supersaturated water were discovered. These waves were significantly correlated with hydropower operation, which was run by hydropeaking (R2=0.82, p<0.001). The TDG saturation varied between 99% and 108%, with a median of 105%. The term "saturopeaking" is introduced for these waves, defined as the artificial, rapid, periodic and frequent fluctuation of gas saturation caused by hydropeaking. Hydropeaking is recognized as hydropower operation that rapidly fluctuates according to the electricity market demand. Though the observed TDG saturation levels were moderate and not likely to cause acute effects on biota, we expect that the observed saturopeaking may have significant ecological impacts in general, especially in cases with TDG saturation levels >110^% which is considered as potentially lethal for fish in rivers.