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Abstract

Anthropogenic barriers such as tidal gates impair animal migration and ecological continuity. For migratory fishes, barriers may alter or impeach both upstream and downstream migrations. Juveniles of European eels, namely glass eels, have to pass those barriers when migrating upstream, whereas they have limited swimming capacities and are dependent on several environmental variables. Hydrological conditions, which can be modified by barrier management, are critical for glass eels. Here, we investigated the links between hydrological conditions and barrier management and the recruitment of glass eels at a connection between the Mediterranean Sea and a lagoon. To do that, we modeled the recruitment of glass eels over ten seasons accounting for temporal variations and co-variable effects. We modeled three aspects of recruitment: presence, level, and composition. We found that monthly and inter-annual effects explained the main part of the variations in glass eel recruitment but accounting for environmental effects improved our models, with a positive effect of temperature, for instance. We associated low levels of catches and recruitment of rather old individuals to high water flow rates when water flows out of the lagoon. Those results call for further studies on how sluice gate management may improve glass eel recruitment and seem to indicate that local stakeholders should adjust the number of open gates depending on the water level on both sides of the barrier.

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The European eel (Anguilla anguilla, Linnaeus 1758) is a critically endangered species that completes its life cycle between the freshwaters of Europe (and North Africa) and spawning grounds in the Sargasso Sea. Its marine larvae are planktonic and metamorphose into post-larval glass eels near the coast. The relationship between environmental variables and the migration of glass eels into brackish water and estuaries is not well understood, but has important implications for European eel conservation and management. Using linear mixed effects (LME) models, we investigated the combined role of meteorological (wind direction and strength and rainfall), oceanographical (tide amplitude), astronomical (moon phase and illumination), temporal and water quality variables on the timing and magnitude of glass eel migration from marine to brackish waters in the Comacchio Lagoon, Northern Adriatic Sea. We used this site as a test case because it was historically prominent in glass eel wild harvest for extensive aquaculture in the Mediterranean region. We found that meteorological factors and tide amplitude had a weak effect on the migration of glass eels into the lagoon. On the contrary, astronomical variables strongly influenced glass eel movement, with waxing phases and intermediate illumination enhancing movement towards the lagoon. We also identified at least two distinct migration pulses during our study period, which were not affected by temporal and water quality variables. Temporal and water quality variables affected body size of glass eels, with bigger glass eels caught later in the season. Identifying the importance of environmental variables affecting glass eels recruitment to brackish waters can help promoting eel stocks restoration through local management.
Article
Tidal barrages on water ways constitute a major threat for diadromous fish species such as the critically endangered European eel (Anguilla anguilla L.). An unobstructed migration route between the spawning area in the Sargasso Sea and the freshwater growth habitats in Europe is crucial for the European eels’ long-term survival. Every spring however, millions of glass eels arrive at the European coast to find their inland migration route blocked. Adjusted tidal barrage management (ATBM), i.e. setting the tidal sluice doors ajar during tidal rise, is one of the mitigation measures to enhance glass eel colonisation. Although this management substantially improves the number of incoming glass eels, the fate of these migrants, confronted with abruptly shifting conditions upon sluice passage, remains un-investigated. Glass eel upstream dispersion after sluice passage by means of ATBM was investigated in a drainage canal connecting a small estuary with an adjacent polder area in Belgium. Large numbers of glass eel caught with eel ladders installed at a pumping station about 800 m upstream the tidal barrage indicate that glass eels were well able to switch to an active counter current swimming behaviour to continue upstream dispersion. On the other hand, some glass eels did not initiate active migration but settled in artificial substrates placed in the canal instead. These residents however appeared very successful in using the canal as feeding ground and in time substantially increased their overall fitness. The appearance of both migration strategies (switch to active migration or successful settlement) in this canal shows the potential of ATBM as a valuable tool for the management and restoration of the European eel population and calls for a wide application of this cost-efficient mitigation measure.
Article
Count data are common in many fields such as public health. Hurdle models have been developed to model count data when the zero count could be either inflated or deflated. However, when data are repeatedly collected over time and spatially correlated, it is very challenging to model the data appropriately. For example, to study health professional shortage areas, the number of primary care physicians along with other demographic characteristics are collected at the county level in the USA and over different years. Since the data are repeatedly collected over time, counties are nested within the state, and adjacent counties are geographically correlated, the dependence structure of the data is very complex. We develop a Bayesian hurdle model with multilayered random effects to incorporate this complex structure. We use a time‐varying random effect for each state to capture the time effect at the state level, and a temporal thin plate spline to capture the spatiotemporal correlation across different counties. We use STAN to obtain samples for inference from the posterior distribution. By using the model proposed, we can identify the important factors which impact health professional shortage areas. Simulation studies also confirm the effectiveness of the model.
Article
L`évolution pigmentaire des civelles est analysée à partir de six échantillons de civelles prélevés en estuaire de Vilaine, et placés dans des cages en estuaire et en eau douce. Les stades pigmentaires des civelles sont lus trois fois par semaine. L`effet de la manipulation sur le vieillissement est contrôlé à l`aide de lots témoins qui sont laissés dans le milieu et analysés en fin d`expérimentation. L'analyse de 9 920 civelles montre que la température est le principal facteur de la cinétique de pigmentation. La durée nécessaire pour passer de 88 à 99 % de jeunes stades (VB et VIA0) à 86 à 99 % de stades plus avancés (VIA2, VIA3 et VIA4) varie de 6 jours à 18 °C à 45 jours à 8°C. La salinité agit comme facteur secondaire en ralentissant la pigmentation. L`anesthésie et la lecture des stades pigmentaires n`affectent pas la pigmentation.
Article
Hatchery-reared salmon smolt used for supplementary stocking often display poor migration behavior compared to wild smolt, which reduces the success of this management action. Oxazepam, an anxiolytic drug, has been shown to intensify salmon smolt migration in mesocosm experiments, and treatment with this drug has, therefore, been suggested as a management option to improve downstream smolt migration. In this study, we tested this by assessing migration performance of hatchery-reared Atlantic salmon (Salmo salar) smolt along a 21-km long natural river-to-sea migration route in a boreal river in Northern Sweden. Using acoustic telemetry, the migration rate and survival of smolt that had been exposed to oxazepam (200 μg L−1, N = 20) was monitored and compared with a control group (N = 20) of unexposed smolt. Exposed smolt took significantly longer time to initiate migration after release compared to the control fish, but after that we observed no significant difference in downstream migration speed. However, exposed smolt had considerably higher probability of being predated on compared to control smolt. We attribute these results to increased risk-taking and higher activity in oxazepam-exposed smolt, which in turn increased initial non-directional exploratory behavior and decreased predator vigilance. These results are discussed based on current concerns for ecological implications of behavioral modifications induced by pharmaceutical pollution and climate change. We conclude that exposure to oxazepam is an unsuitable management option to prime migration of reared salmon in natural systems.
Article
This chapter provides information on nomenclature and systematics, description, distribution, ecology, food and eating behavior, reproduction and reproductive behavior, economic importance, protection status, and conservation of Anguilla. Species of the genus Anguilla are widely distributed in most tropical, subtropical and temperate areas of the world. They are present on all continents, except in Antarctica. Juvenile (yellow, green) and subadult (silver) eels have a very elongated, serpentiform, circular‐shaped body, which is slightly compressed particularly close to the caudal end. Within its wide range, the eel visits coastal waters (coastal areas, estuaries, lagoons) as well as continental waters (rivers, lakes, ponds, marshes, reservoirs). Transparent eels progressively become pigmented because of the development of cells enriched with melanic pigment (black), and thus become pigmented glass eels. The eel is generally the most abundant species in Mediterranean lagoons, especially in the Adriatic and in the northern sector of the western basin.
Article
Connectivity between freshwater habitats and marine areas is heavily obstructed by anthropogenic structures (e.g. weirs, pumping stations, sluices…), leading to a high pressure on diadromous fish populations. A better understanding of fish migration behaviour in relation to these barriers is needed to take proper mitigation actions. We investigated the impact of migration barriers on downstream migrating European eel (Anguilla anguilla L.) by tracking 50 acoustically tagged eels between July 2012 and March 2015 in a Belgian polder area. The study area was selected due to the presence of a wide range of migration barriers, such as two pumping stations, a weir and tidal sluices. These structures regulate the water level, resulting in discontinuous flow conditions. The results showed that migration was primarily nocturnal and discharge appeared to be the main trigger for migration in the polder. We also observed substantial delays and exploratory behaviour near barriers. Delays can have a serious impact on eels since their energy resources are limited for a successful trans-Atlantic migration. In addition, delays and exploratory behaviour can also increase predation and disease risk. The obtained knowledge can contribute to efficient management such as improved fish passage and guidance solutions.
Article
The relationships between the migratory behavior, methylmercury (MeHg) concentrations, oxidative stress response and detoxification processes were investigated in glass eels collected in marine (Molliets) and estuarine (Urt) waters (Adour estuary, South West France) at the end of the fishing season (April). Glass eel migratory behavior was investigated in an experimental flume according to their response to dusk. Fish responding to the decrease in light intensity by ascending in the water column and moving with or against the flow were considered as having a high propensity to migrate (migrant). Glass eels still sheltering at the end of the 24 h catching period were considered as having a low propensity to migrate and were called non-migrant. Our results provide some evidence that estuarine glass eels were bigger, presented a higher propensity to migrate and a lower oxidative stress response than marine glass eels. This might reflect a selection process, some marine glass eels progressively settling or dying before reaching Urt and/or a change in feeding behavior. In April, glass eels restart feeding in the Adour estuary which might decrease the oxidative stress possibly related to starvation, and enhance migration. MeHg concentrations was significantly higher in non-migrant than in migrant glass eels and it is suggested that non-migrant glass eels might present a higher vulnerability to stress (at least contamination and/or starvation), although the underlying mechanisms remain to be elucidated.
Article
Passage of fish through hydropower dams is associated with mortality, delay, increased energy expenditure and migratory failure for migrating fish and the need for remedial measures for both upstream and downstream migration is widely recognised. A functional fish passage must ensure safe and timely passage routes that a substantial portion of migrating fish will use. Passage solutions must address not only the number or percentage of fish that successfully pass a barrier, but also the time it takes to pass. Here, we used radiotelemetry to study the functionality of a fish bypass for downstream-migrating wild-caught and hatchery-released Atlantic salmon smolts. We used time-to-event analysis to model the influence of fish characteristics and environmental variables on the rates of a series of events associated with dam passage. Among the modelled events were approach rate to the bypass entry zone, retention rates in both the forebay and the entry zone and passage rates. Despite repeated attempts, only 65% of the tagged fish present in the forebay passed the dam. Fish passed via the bypass (33%), via spill (18%) and via turbines (15%). Discharge was positively related to approach, passage and retention rates. We did not detect any differences between wild and hatchery fish. Even though individual fish visited the forebay and the entry zone on multiple occasions, most fish passed during the first exposures to these zones. This study underscores the importance of timeliness to passage success and the usefulness of time-to-event analysis for understanding factors governing passage performance.
Article
Analysis of glass eels catches from 1927 to 1998 in the Adour estuary showed different periods of successive decrease. Fishing periods between 1985 to 1993 were studied in order to observe impact of hydroclimatic conditions on the level of catch per unit of effort. This study took place in January when glass eels accessibility by the fishery was generally the highest. With hand and trawl netting that filtered the upper part of the water column, only individuals located close to the surface were accessible. Consequently, factors acting on the migration of glass eels in the water column have a great importance on the fishing success. Among environmental conditions, moon phase and temperature influence were determinant factors on catch variability but a lack of relationship between tide coefficient and catches is found. These studies showed the importance of night brightness on the migratory behaviour of glass eels and consequently on the accessibility of that species by the fishery.
Article
Given the importance of reliable recruitment estimates when assessing temperate eel stocks and enforcing appropriate management measures, surprisingly few analytical tools have been developed to estimate yearly glass eel recruitment. Of the models that do exist, large-scale models generally rely on strong assumptions relating to fishing activity, while other models generally estimate recruitment at the river basin scale. With the aim of filling this gap, we developed the GEREM (Glass Eel Recruitment Estimation Model) to estimate glass eel recruitment at different nested spatial scales. Our model simultaneously estimates annual recruitment at the river catchment level, at an intermediate spatial scale such as Eel Management Units (EMUs), and at a larger scale (e.g., a country). Provided sufficient data become available in the future, the analysis could be extended to the scale of the distribution area, which would be consistent with the population scale. In this study, the model was applied to France, using various recruitment indices obtained from 1960 to 2013. This provided trends and absolute recruitment estimates consistent with current expert knowledge. A sensitivity analysis was carried out to assess the robustness of results to sources of uncertainty. This type of model fills an important gap in the range of quantitative tools presently available to estimate recruitment. It could be used in the future to establish total allowable catches in countries such as France where glass eels are fished commercially.
Article
Juvenile Chinook Salmon Oncorhynchus tshawytscha emigrating from natal tributaries of the Sacramento River, California, must negotiate the Sacramento-San Joaquin River Delta (hereafter, the Delta), a complex network of natural and man-made channels linking the Sacramento River with San Francisco Bay. Fish that enter the interior and southern Delta—the region to the south of the Sacramento River where water pumping stations are located—survive at a lower rate than fish that use alternative migration routes. Consequently, total survival decreases as the fraction of the population entering the interior Delta increases, thus spurring management actions to reduce the proportion of fish that are entrained into the interior Delta. To better inform management actions, we modeled entrainment probability as a function of hydrodynamic variables. We fitted alternative entrainment models to telemetry data that identified when tagged fish in the Sacramento River entered two river channels leading to the interior Delta (Georgiana Slough and the gated Delta Cross Channel). We found that the probability of entrainment into the interior Delta through both channels depended strongly on the river flow and tidal stage at the time of fish arrival at the river junction. Fish that arrived during ebb tides had a low entrainment probability, whereas fish that arrived during flood tides (i.e., when the river's flow was reversed) had a high probability of entering the interior Delta. We coupled our entrainment model with a flow simulation model to evaluate the effect of nighttime closures of the Delta Cross Channel gates on the daily probability of fish entrainment into the interior Delta. Relative to 24-h gate closures, nighttime closures increased daily entrainment probability by 3 percentage points on average if fish arrived at the river junction uniformly throughout the day and by only 1.3 percentage points if 85% of fish arrived at night. We illustrate how our model can be used to evaluate the effects of alternative water management actions on fish entrainment into the interior Delta.
Article
As part of flood protection and land reclamation schemes, tide gates allow rivers to discharge to sea when open, and prevent salt water intrusion when closed. Their impact on diadromous fish migration between essential spawning and rearing habitats, and the effectiveness of mitigation measures, have received little consideration. The River Meon, UK, discharges to sea through four top-hung counterbalanced tide gates. In March 2012, the gates were replaced with new ones of the same design, but with an orifice installed in two of them partly to improve fish passage. Sixty downstream migrating juvenile sea trout, Salmo trutta, were trapped approximately 4.9 km upstream of the tidal limit and tagged with acoustic transmitters in April 2011 (n= 30) and 2012 (n = 30). Tagged individuals were detected by acoustic receivers placed near the tide gates before (year 1) and after (year 2) orifice installation. Of the fish that approached the tide gates, 95.8% and 100.0% successfully passed in years 1 and 2, respectively. The speed of migration at the gates was slower than for upstream and downstream reaches, and was positively related to percentage of time the gates were open. Presence of the orifices did not influence delay. Overall, top-hung tide gates delayed sea trout migration, potentially increasing the risk of predation and energy expenditure during the vulnerable juvenile life stage.
Article
The aim of this review is to present an overview of the sex differentiation and sex determination processes in eels in relation to the urgent need to provide scientific knowledge to better protect and manage the Anguilla genus. Indeed, the global decline of the three main temperate eel stocks, Anguilla anguilla, Anguillidae (Fisheries Management and Ecology, 2003, 10, 365); Anguilla japonica, Anguillidae (Casselman, Eel Biology, Springer Japan, 2003, 293) and Anguilla rostrata, Anguillidae (Tatsukawa, Eel Biology, Springer, Japan, 2003, 255), raises concerns about the necessity to better understand all stages of the life cycle of eels (Righton and Walker, Journal of Fish Biology, 2013, 83, 754). Little is known about the mechanisms involved in the production of males and females in this species with environmental sex determination. Previous reviews identifying the density of individuals as the major factor influencing sex determination were undertaken (Krueger and Oliveira, Environmental Biology of Fishes, 1999, 55, 381; Reviews in Fish Biology and Fisheries, 2005, 15, 37). Here, we review the current advances on the subject, focusing on the roles of early growth rate and interindividual relationships, which are mechanisms underpinned by density, as well as the sex differentiation process, and we question how this knowledge might influence global conservation measures.
Article
Tide gates form a temporal barrier to fish migration, closing during the flood tide and opening during the ebb, primarily for flood prevention and land reclamation. Their impact on downstream adult migration of the critically endangered European eel, Anguilla anguilla, is unknown. The River Stiffkey, UK, has three top-hung tide gates (one counterbalanced, two not) through which it discharges into the North Sea. Adult eels of silver appearance (n = 118) were caught between 0.5 and 6.0 km upstream from the tide gates in Autumn 2011 and implanted with 23 mm half-duplex passive integrated transponder (PIT) tags. Tagged individuals were detected by PIT antennae located near the tide gates. Of the eels tagged, 80 were detected actively migrating downstream to the gates. Escapement past the gates was 98.3 %. Speed of migration was slower near the gates than for an unimpeded upstream reach and was positively and negatively related to mean degree of gate opening and mean light intensity, respectively. When the largest gate was modified through installation of an orifice intended to improve upstream passage of sea trout and juvenile eels, downstream migration was more rapid when it was operating. However, video analysis revealed that eels did not pass through the orifice, meaning that faster migration may have been a result of the gates being open on more occasions when eels initially approached them, or the lower tides and upstream saline intrusion that occurred during these periods. Top-hung tide gates in the River Stiffkey delayed eel migration, potentially increasing the risk of predation and energy expenditure immediately prior to a 5000–6000 km migration to spawning grounds in the Sargasso Sea.
Article
Globally, diadromous species are at risk from fragmentation by damming of rivers, and a host of other anthropogenic factors. On the United States Atlantic Coast, where diadromous fish populations have undergone dramatic declines, restoration programs based on fishway construction and hatcheries have sustained remnant populations, but large-scale restoration has not been achieved. We examine anadromous fish restoration programs on three large Atlantic Coast rivers, the Susquehanna, Connecticut, and Merrimack with multiple mainstem hydropower dams, most with relatively low generating capacity. Mean passage efficiencies through fishways on these rivers from the first dam to the spawning grounds for American shad are less than 3%. The result is that only small fractions of targeted fish species are able to complete migrations. It may be time to admit failure of fish passage and hatchery-based restoration programs and acknowledge that significant diadromous species restoration is not possible without dam removals. The approach being employed on the Penobscot River, where dams are being removed or provided the opportunity to increase power generation within a plan to provide increased access to habitat, offers a good model for restoration. Dammed Atlantic Coastal rivers offer a cautionary tale for developing nations intent on hydropower development, suggesting that lasting ecosystem-wide impacts cannot be compensated for through fish passage and hatchery technology.
Article
In an attempt to restore the connectivity of fragmented river habitats, a variety of passage facilities have been installed at river barriers. Despite the cost of building these structures, there has been no quantitative evaluation of their overall success at restoring fish passage. We reviewed articles from 1960 to 2011, extracted data from 65 papers on fish passage efficiency, size and species of fish, and fishway characteristics to determine the best predictors of fishway efficiency. Because data were scarce for fishes other than salmonids (order Salmoniformes), we combined data for all non‐salmonids for our analysis. On average, downstream passage efficiency was 68.5%, slightly higher than upstream passage efficiency of 41.7%, and neither differed across the geographical regions of study. Salmonids were more successful than non‐salmonids in passing upstream (61.7 vs. 21.1%) and downstream (74.6 vs. 39.6%) through fish passage facilities. Passage efficiency differed significantly between types of fishways; pool and weir, pool and slot and natural fishways had the highest efficiencies, whereas Denil and fish locks/elevators had the lowest. Upstream passage efficiency decreased significantly with fishway slope, but increased with fishway length, and water velocity. An information‐theoretic analysis indicated that the best predictors of fish passage efficiency were order of fish (i.e. salmonids > non‐salmonids), type of fishway and length of fishway. Overall, the low efficiency of passage facilities indicated that most need to be improved to sufficiently mitigate habitat fragmentation for the complete fish community across a range of environmental conditions.
Article
Estuarine barriers may significantly reduce the upstream migration of diadromous fish species like the European eel (Anguilla anguilla L.). Previous research showed that limited barrier opening during tidal rise was a cost-efficient and effective mitigation option to improve upstream glass eel migration, without significant intrusion of sea water. This paper aims to optimise this adjusted barrier management to improve eel passage at a tidal barrier complex at the mouth of the River Yser, Flanders, Belgium, one of the most important migration routes for glass eel in Flanders. Specifically, three hypotheses were tested. The first hypothesis analysed the impact of the number of barriers opened on the upstream glass eel migration. The second hypothesis evaluated the relation between the size of the barrier opening and glass eel migration. Finally, we tested whether the suggested adjusted barrier management may lead to a significant increase in conductivity in the River Yser. Increased opening of one barrier appeared more efficient than opening several barriers slightly. Conductivity increased during periods of extreme drought and at base flow, but decreased within 24 h after the first peak flow. This indicates that adjusted barrier management does not entail salt intrusion in the Yser basin, as long as this management is not applied in extremely dry periods. Since the adjusted barrier management is easily implemented and could be applied on numerous tidal barriers, the presented results may contribute to restoration of eel populations worldwide and be of interest to a wide range of river managers and stakeholders.
Article
Fluvial disconnectivity can have important impacts on fish populations, including hindering movement between habitats required for different ontogenic stages. Recruitment of the European eel (Anguilla anguilla) has reduced by over 90% since the early 1980's, in part due to the effect of riverine barriers on its catadromous migration. There is a legislative requirement to restore free passage, increase habitat availability, and limit anthropogenic losses at intakes to aid eel recovery and good ecological status; necessitating an improved understanding of underlying processes. Escapement, route choice, delay at structures, and entrainment at water abstraction points of downstream migrating silver eels were examined using acoustic and Passive Integrated Transponder (PIT) telemetry in the heavily regulated lower river Stour, UK. Downstream migrating adult eel (n = 69) were trapped approximately 10 km upstream of the tidal limit, surgically implanted with an acoustic transducer and PIT transponder, and released between October and December in 2009 and 2010. Movements of tagged individuals were monitored by a linear array of 19 fixed acoustic receivers extending from the release site, through the last 9.2 km of the freshwater catchment. Three groups of water control structures, two water abstraction intakes and several possible routes of migration are present in the reach. Seventy six and 65% of tagged eels escaped from the study reach in 2009 and 2010, respectively. Entrainment at a single intake was the principal cause of loss and positively related to rapid increases in abstraction whilst eels were in the vicinity of the intake. Route choice into the estuary was dependent on discharge over a large intertidal weir; opening regimes of a tidal gate at the termination of the alternative channel; and abstraction rate at a nearby water intake. Long delays (up to 68.5 days) and recurrent behaviour were associated with several structures in the study reach; high variability between individuals reflected the management of spill at weirs. Potential scenarios for minimising entrainment and delay through integrated management of water level control structures and abstraction rates are discussed.
Article
Most fish populations are declining worldwide and their management would benefit from a better estimation of recruitment. In glass eels, field studies suggest that estuarine migratory glass eels are sensitive enough to light to change their vertical location according to factors such as water turbidity and/or moon brightness. The response of glass eel (Anguilla anguilla L.) to light was tested in the laboratory using boxes where fish could choose between a lit and an unlit side. Responses were quantified as the proportion of glass eels remaining in the unlit chamber. Decreasing light levels were used and tested on different “age” glass eels (“age” in days since capture). In addition, measures of light at different depths of the water column were carried out in the Adour estuary (43°30′ N, 1°30′ W). The glass eel light avoidance level was lower in non-pigmented glass eel (less than 10−10 W cm−2), than in pigmented ones (10−9–10−8 W cm−2). These results and field data on the measurement of light energy in the water column of Adour estuary are compared with previously published data on the estuarine migration of glass eel.
Article
The pigmentation development process of glass eels Anguilla anguilla from stage VB to VIA3 was modelled by gamma cumulative functions. These functions varied with respect to the factors temperature and salinity whose effects were adjusted by beta functions. Temperature was shown to accelerate pigmentation, while salinity acted as a secondary factor slowing down the pigmentation. The model fits the development of 15 samples kept at various temperatures and salinities in the Vilaine River, as well as samples monitored at other dates and places in Europe. It allows the prediction of the duration of estuarine residency for glass eels, in winter and spring, in the Atlantic estuaries.
Article
Pollution from a base metal mine on a tributary of the Northwest Miramichi River caused many adult Atlantic salmon, which were on their normal upstream spawning migration, to return prematurely downstream through a counting fence on that river during summer and early autumn. These observations gave an opportunity to document avoidance reactions of salmon to pollution, which has seldom been done in the fishes' natural environment. Downstream returns of salmon rose from between 1 and 3 per cent during 6 years before pollution to between 10 and 22 per cent during 4 years of pollution. Early runs (June–July) of salmon to the headwaters were delayed and reduced in number. Chemical analyses of river water showed levels of Cu2+ and Zn2+ which varied with rates of river discharge. During some periods Cu2+ + Zn2+ concentrations exceeded lethal levels for immature salmon, as established in another (laboratory) study. The threshold concentration for 50 per cent survival of fish under specified temperature conditions is designated as 1·0 toxic unit. Adult salmon in nature showed avoidance reactions at about 0·35–0·43 toxic unit of Cu2+ + Zn2+. A level of 0·8 toxic unit may have blocked all upstream movement. Of the salmon returning downstream because of pollution, about 31 per cent reascended, 62 per cent were not seen again and 7 per cent were taken by angling and commercial fishing below the counting fence. Estimated losses from the stock available in the upper part of the river from 1960 to 1963 varied from 8 to 15 per cent of the total run. There is no evidence that successive year-classes of salmon are growing accustomed to the pollution.