Factors affecting the within-river spawning migration of Atlantic salmon,with emphasis on human impacts

We review factors affecting the within-river 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 within-river 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.     

Migratory behaviour of adult pikeperch (Stizostedion lucioperca) in a lowland river

The behaviour of radio-tagged adult pikeperch (Stizostedion lucioperca (L.)) from two areas in the Danish River Gudenaa were recorded prior to, and during the spawning period. Eight of 13 tagged fish in the lower reaches of the river were located throughout the whole study. Five of these fish moved upstream to various sites in the river prior to spawning, which occurred from late April to June. The three remaining fish moved to the fjord. These movements were interpreted as a spawning migration, and it is suggested that the lower reaches of the River Gudenaa constitute an over-wintering area for pikeperch, which use different spawning areas. Ten pikeperch caught just downstream of an impassable hydropower plant were radio-tagged and translocated upstream of the dam to a reservoir. Within a week, half of the fish moved to a lake situated more than 30 km upstream the reservoir. This behaviour is hypothesised to be a homing response. The study reveals that the pikeperch is a highly mobile species with a complicated migration pattern, even in relatively small river systems.     

Upstream passage problems for wild Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) in a regulated river and its effect on the population

Due to hydropower development, the upstream migration of wild anadromous salmon and brown trout is impaired in many European rivers, causing negative effects on the long-term survival of natural salmonid populations. This study identified problems for Atlantic salmon during upstream migration in a regulated river in northern Sweden, Umeälven (mean flow: 430 m³ s^?1). Tagging from 1995 to 2005 involved radio tags (n = 503), PIT tags (n = 1574) and Carlin tags (n = 573) to study the spawning migration of salmon from the coast past the regulated section of the river to a fish ladder at the dam/spillway 32 km upriver. The results demonstrate that migration success from the coast to the fish ladder varied between 0% and 47% among years, indicating an average loss of 70% of potential spawners. Discharge from the turbines attracted the salmon away from the bypass route. Echo-sounding in the turbine outlet showed that salmon were normally found at 1–4 m depths. They responded with upstream and/or downstream movements depending on flow changes; increased spill in the bypass channel attracted salmon to the bypass. Once in the bypass channel, salmon could be delayed and had difficulties passing the first rapid at high spills. Additional hindrances to upstream migration were found at rapids and the area of the fish ladder, located further upstream in the regulated river section. The average migration duration was 44 days from the estuary to the top of the fish ladder, with large variation among individuals within years. Modelling the salmon population dynamics showed a potential population increase of 500% in 10 years if the overall migration success could be improved from the current 30% to levels near 75%. Consequently improved migration facilities at the regulated river section should be implemented to achieve a long-term sustainability of these threatened anadromous salmonids.     

Dynamic Modeling and Experiment of a Fish Robot with a Flexible Tail Fin

This paper presents the dynamic modeling of a flexible tail for a robotic fish. For this purpose firstly, the flexible tail was simplified as a slewing beam actuated by a driving moment. The governing equation of the flexible tail was derived by using the Euler-Bernoulli theory. In this equation, the resistive forces were estimated as a term analogous to viscous damping. Then, the modal analysis method was applied in order to derive an analytical solution of the governing equation, by which the relationship between the driving moment and the lateral movement of the flexible tail was described. Finally, simulations and experiments were carried out and the results were compared to verify the accuracy of the dynamic model. It was proved that the dynamic model of a fish robot with a flexible tail fin well explains the real behavior of robotic fish in underwater environment.     

An optimal stopping approach for onset of fish migration

Comprehending life history of migratory fish, onset of migration in particular, is a key biological and ecological research topic that still has not been clarified. In this paper, we propose a simple mathematical model for the onset of fish migration in the context of a stochastic optimal stopping theory, which is a new attempt to our knowledge. Finding the criteria of the onset of migration reduces to solving a variational inequality of a degenerate elliptic type. As a first step of the new mathematical modeling, mathematical and numerical analyses with particular emphasis on whether the model is consistent with the past observation results of fish migration are examined, demonstrating reasonable agreement between the theory and observation results. The present mathematical model thus potentially serves as a simple basis for analyzing onset of fish migration.     

Analytical solution for a hybrid Logistic-Monod cell growth model in batch and continuous stirred tank reactor culture

Monod and Logistic growth models have been widely used as basic equations to describe cell growth in bioprocess engineering. In the case of the Monod equation, the specific growth rate is governed by a limiting nutrient, with the mathematical form similar to the Michaelis–Menten equation. In the case of the Logistic equation, the specific growth rate is determined by the carrying capacity of the system, which could be growth-inhibiting factors (i.e., toxic chemical accumulation) other than the nutrient level. Both equations have been found valuable to guide us build unstructured kinetic models to analyze the fermentation process and understand cell physiology. In this work, we present a hybrid Logistic-Monod growth model, which accounts for multiple growth-dependent factors including both the limiting nutrient and the carrying capacity of the system. Coupled with substrate consumption and yield coefficient, we present the analytical solutions for this hybrid Logistic-Monod model in both batch and continuous stirred tank reactor (CSTR) culture. Under high biomass yield (Y_x/s) conditions, the analytical solution for this hybrid model is approaching to the Logistic equation; under low biomass yield condition, the analytical solution for this hybrid model converges to the Monod equation. This hybrid Logistic-Monod equation represents the cell growth transition from substrate-limiting condition to growth-inhibiting condition, which could be adopted to accurately describe the multi-phases of cell growth and may facilitate kinetic model construction, bioprocess optimization, and scale-up in industrial biotechnology.     

Using "travel time" data to characterize the behavior of migrating animals

For migratory species, duration of migration, or "travel time," is often a critical variable in determining the cost of migration. Observed travel times are the result of both environmental factors such as air or water currents and the behavior of individuals. In an effort to distinguish among these components, I developed a migration model based on an advection-diffusion equation that characterizes population movements in terms of two biologically meaningful parameters: migration rate and rate of population spread. I applied the model to travel time data from juvenile chinook salmon (Onchorhynchus tshawytscha), which were tagged during their seaward migration. The tagged fish originated from three separate evolutionarily significant units (ESUs) as classified by the U. S. National Marine Fisheries Service. The model was expanded by allowing migration and diffusion rates to vary with fish length and river flow. Variability in travel times explained by these factors was strikingly similar from year to year within ESUs, and the migratory behavior revealed by the analysis was consistent with the life-history patterns that distinguish the ESUs. The approach presented here is easily adaptable to a wide range of migratory species and may be particularly useful for predicting how at-risk populations respond to variable conditions in regulated or otherwise disturbed migration habitats.     

Tidal and diel timing of river entry by adult Atlantic salmon returning to the Aberdeenshire Dee, Scotland

The timing of salmon migration from the estuary of the Aberdeenshire Dee into the river in relation to tidal phase and time of day was studied by combined acoustic and radio-tracking of individual fish and by analysing records of untagged fish from a resistivity fish counter 0.8 km upstream from the tidal limit. Up-estuary movements that led to river entry were predominantly nocturnal and tended to occur during the ebb tide. Penetration into the non-tidal reaches of the river also tended to occur at night, but the timing of salmon movements was no longer significantly associated with tidal phase. The tracking data suggested that the reduction in the strength of the association between salmon migration and tidal phase resulted from variability in rates of progress from the estuary to the river. This variability may have been random, or related to changes in migratory behaviour during entry to fresh water. A simulation of upstream progress by groups of salmon illustrated the decay of the relationship between salmon movements and tidal phase and the advance of the average tidal phase of observed salmon movements with increasing distance of the observation point from where movements were initiated. The magnitude of these effects depended on the average rate of upstream progress and variability in the rate of progress, being greatest when upstream progress was slow and variable. These results highlight a limitation of point observations of migration with regard to identifying environmental stimuli for migration and quantifying their effects.     

Variation in local abundance and species richness of stream fishes in relation to dispersal barriers: implications for management and conservation

1. Barriers to immigration, all else being equal, should in principle depress local abundance and reduce local species richness. These issues are particularly relevant to stream‐dwelling species when improperly designed road crossings act as barriers to migration with potential impacts on the viability of upstream populations. However, because abundance and richness are highly spatially and temporally heterogeneous and the relative importance of immigration on demography is uncertain, population‐ and community‐level effects can be difficult to detect. 2. In this study, we tested the effects of potential barriers to upstream movements on the local abundance and species richness of a diverse assemblage of resident stream fishes in the Monongahela National Forest, West Virginia, U.S.A. Fishes were sampled using simple standard techniques above‐ and below road crossings that were either likely or unlikely to be barriers to upstream fish movements (based on physical dimensions of the crossing). We predicted that abundance of resident fishes would be lower in the upstream sections of streams with predicted impassable barriers, that the strength of the effect would vary among species and that variable effects on abundance would translate into lower species richness. 3. Supporting these predictions, the statistical model that best accounted for variation in abundance and species richness included a significant interaction between location (upstream or downstream of crossing) and type (passable or impassable crossing). Stream sections located above predicated impassable culverts had fewer than half the number of species and less than half the total fish abundance, while stream sections above and below passable culverts had essentially equivalent richness and abundance. 4. Our results are consistent with the importance of immigration and population connectivity to local abundance and species richness of stream fishes. In turn, these results suggest that when measured at appropriate scales (multiple streams within catchments), with simple protocols amenable to use by management agencies, differences in local abundance and species richness may serve as indicators of the extent to which road crossings are barriers to fish movement and help determine whether road‐crossing improvements have restored connectivity to stream fish populations and communities.     

Survival of captured and relocated adult spring-run Chinook salmon Oncorhynchus tshawytscha in a Sacramento River tributary after cessation of migration

We studied the efficacy of the process for capture and upstream relocation of 26 adult spring-run Chinook salmon in Butte Creek, California in 2009. These fish had ceased volitional upstream migration prior to reaching their summer holding habitat. The purpose of the relocation was to move fish upstream of two water diversion dams and release them in a part of the stream from which they could presumably swim to cool summer holding habitat, then spawn in the fall. Fish were netted, transported by truck, given an esophageal radio tag/temperature tag, and released. Radio tagging proved to be a useful technique for determining the survival and movement of relocated fish and temperature tags provide useful information to determine thermal exposure and time of death. Twenty-three tags (88 %) were recovered, compared with a 10 % tag recovery rate for an earlier study using fin clips. Most tags were recovered within 3.5 km upstream and 1 km downstream of the release site. A single tag was recovered 6 km upstream. No fish were determined to have survived to spawn. Temperature tag data indicate that most of the salmon died within 2–6 days after the relocation operation. After preventative measures have been exhausted, future relocations efforts, in any setting, should consider (1) intervention as soon as fish cease volitional migration but before they are exposed to further deleterious conditions (2) monitoring environmental conditions to choose appropriate release sites (3) evaluation of disease transmission risk, and (4) handling practices that minimize potential stress due to air immersion and thermal shock.     

Sequential fishways reconnect a coastal river reflecting restored migratory pathways for an entire fish community

River regulation infrastructure has been implicated in worldwide aquatic biodiversity loss. In‐stream barriers such as weirs prevent fish migration and the impact can be particularly severe for diadromous species. Fishways are frequently installed on in‐stream barriers to reconnect migratory pathways and rehabilitate diadromous fish populations. Sequential fishways may be necessary in rivers where multiple barriers prevent migration. We monitored a coastal fish community's response to fish passage restoration at 10 predominantly low‐level weirs in the freshwater reaches of the Nepean River in southeastern Australia. Few studies have simultaneously assessed fish community passage at successive fishways within a river system. Prior to the installation of the fishways, there was a gradient of reduced species diversity in an upstream direction including the absence of many diadromous species, despite the regular inundation frequency of most weirs. Following the installation of the fishways, species diversity was still greater in the downstream monitoring sites; however, there was evidence of a positive change in fish community structure in middle and upstream sites. Most notably, three diadromous species rapidly expanded their distribution upstream and one amphidromous species expanded its downstream distribution. This study demonstrates appropriately designed successive fishways can successfully reconnect river systems for an entire fish community, encompassing species with a broad range of swimming abilities and diverse life histories.     

Quantitative methods for analysing cumulative effects on fish migration success: a review

It is often recognized, but seldom addressed, that a quantitative assessment of the cumulative effects, both additive and non-additive, of multiple stressors on fish survival would provide a more realistic representation of the factors that influence fish migration. This review presents a compilation of analytical methods applied to a well-studied fish migration, a more general review of quantitative multivariable methods, and a synthesis on how to apply new analytical techniques in fish migration studies. A compilation of adult migration papers from Fraser River sockeye salmon Oncorhynchus nerka revealed a limited number of multivariable methods being applied and the sub-optimal reliance on univariable methods for multivariable problems. The literature review of fisheries science, general biology and medicine identified a large number of alternative methods for dealing with cumulative effects, with a limited number of techniques being used in fish migration studies. An evaluation of the different methods revealed that certain classes of multivariable analyses will probably prove useful in future assessments of cumulative effects on fish migration. This overview and evaluation of quantitative methods gathered from the disparate fields should serve as a primer for anyone seeking to quantify cumulative effects on fish migration survival.     

Sources of fish in the ephemeral western iishana region of the Cuvelai–Etosha Basin in Angola and Namibia

The triangle between the Kavango and Kunene rivers is drained by the Cuvelai, an ephemeral and deltaic drainage system covering more than 100 000 km². In good rainfall years, the area becomes populated by fish communities dominated by five species migrating southward towards the endorheic Etosha Pan, the basin’s terminal sump. When water dries up, fish subsequently die-off and their sudden appearance in rainy years has captivated scientists for decades. The current study was prompted by hitherto untapped indigenous knowledge through narratives of opportunistic fish harvesting of migrating fish at temporary connections between the Kunene River and the Cuvelai- Etosha Basin. A reconnaissance fish survey in 2017 was complemented by digital satellite images and elevation data analyses. Results support the presence of at least three major ephemeral fish migration routes. The dominant fish genera migrating upstream in Kunene tributaries comprise Enteromius, Oreochromis and Clarias, all eurytopic and known to undertake upstream, lateral and downstream migrations on floodplains. Although other notable fish refugia in the Cuvelai–Etosha Basin are yet to be identified, there is a necessity for the protection and management of these migration routes in tandem with studies on the nature and extent of this inter-basin fish migration under climate change and variability.     

Thermal biology and bioenergetics of different upriver migration strategies in a stock of summer-run Chinook salmon

By combining biotelemetry with animal-borne thermal loggers, we re-created the thermal histories of 21 summer-run Chinook salmon (Oncorhynchus tshawytscha) migrating in the Puntledge River, a hydropower impacted river system on Vancouver Island, British Columbia, Canada. Daily maximum water temperatures in the Puntledge River during the summer-run adult Chinook salmon migration and residency period frequently exceeded 21 °C, a value that has been observed to elicit behavioral thermoregulation in other Chinook salmon populations. We therefore compared river temperatures to body temperatures of 16 fish that migrated through the river to understand if cool-water refuge was available and being used by migrants. In addition, we used thermal histories from fish and thermal loggers distributed in the river to model the effect of thermal habitat on energy density using a bioenergetics model. In general, we found no evidence that cool-water refuge existed in the river, suggesting that there is no opportunity for fish to behaviorally thermoregulate during upriver migration through the regulated portion of the river. Of the thermal histories used in the bioenergetics model, fish that reached an upstream lake were able to access cooler, deeper waters, which would have reduced energy consumption compared to fish that only spent time in the warmer river. Consequently, the Puntledge River water temperatures are likely approaching and in some cases exceeding the thermal limits of the summer-run Chinook salmon during the spawning migration. Further warming may cause more declines in the stock.     

Pre-spawning migration of adult Pacific lamprey, <Emphasis Type="BoldItalic">Entosphenus tridentatus</Emphasis>, in the Willamette River,Oregon, U.S.A.

We describe the migration distances and timing of the adult Pacific lamprey, Entosphenus tridentatus, in the Willamette River Basin (Oregon, U.S.A.). We conducted aerial surveys to track radio-tagged fish upstream of a major waterfall and hydropower complex en route to spawning areas. We detected 24 out of the 43 fish that passed the waterfall-hydropower complex. Of the detected fish, 17 were detected multiple times. Their maximum migration distance upstream in the mainstem Willamette approximated a normal distribution. The maximum distance migrated upstream did not significantly correlate with total body length (r = −0.186, P = 0.385) or date that the fish passed Willamette Falls (r = −0.118, P = 0.582). Fish migrated primarily during the spring to early summer period before stopping during the summer, when peak river temperatures (≥20°C). However, at least three fish continued to migrate upstream after September. Behavior ranged from relatively slow migration, followed by holding; to rapid migration, followed by slow migration further up in the basin. This study provides a basis for informing more detailed research on Pacific lamprey in the future.     

Fish ladders select fish traits on migration–still a growing problem for natural fish populations

We investigated the potential selective effect of fish ladders on physiological and morphological profiles of the curimbatá, Prochilodus lineatus, during reproductive migration in Brazil. We registered sex, body weight and length, plasma glucose, hepatosomatic and gonadosomatic indices (HSI and GSI, respectively), hematocrit, leucocrit, blood cell and nucleus areas, and the diameter of white and red muscle fibers in fish sampled at the bottom (downstream) and at the top (upstream) of a fish ladder at a hydroelectric dam. Males and females at the top of the ladder showed higher size (weight and length), white muscle fiber diameters, plasma glucose levels and lower hematocrit when compared with those at the bottom. These size and muscle traits assist fish to overcome the ladder barrier and bypass the dam, an effort that might be reflected in the glucose levels. Females also showed higher GSI at the top of the fish ladder, a trait possibly facilitating their reproduction upstream. These results indicate that a dam system favors fish with specific morphological–physiological profile. This may have a strong influence upon upstream fish populations over generations and implies the presence of artificial selective pressure.     

Annual movement and migration of adult pikeperch in a lowland river

The downstream migration of 46–66 cm radio-tagged adult pikeperch Stizostedion lucioperca , in the River Gudenaa, Denmark, from late September to late December, coincided with a reduction in the frequent occurrence of prey fish in the tagging area, and hence was interpreted as a feeding migration. The upstream migration from March to mid-April in spring took place prior to spawning which occurred from late April to June.     

Upstream migration of adult Atlantic salmon past a fish counter weir in the Aberdeenshire Dee, Scotland

During their upstream migration through the lower reaches of the Aberdeenshire Dee, Scotland, radio-tagged adult Atlantic salmon passed a Crump weir-based fish counter without substantial delay over a range of river flows and water temperatures. Tagged salmon were detected sequentially by automatic listening stations placed along the river and were detected at the counter site for a median time of 18 min, compared with a median time of 5 min before the weir was built. One fish out of 16 spent 6·7 h in the vicinity of the weir, but such migratory pauses also occurred before the weir existed. The results suggest that the number of salmon crossing the counter per unit time is a function of the number of fish present downstream and their tendency to migrate upstream, without being limited by their ability to surmount the weir. The apparent slight delay at the weir is probably insignificant in terms of the overall progress of riverine migration and in the context of using counter records to relate salmon movements to environmental Variables.     

Computational and mathematical modeling of the effects of tailbeat frequency and flexural stiffness in swimming fish

In this paper we describe how we combine computational and mathematical models to form virtual fish to explore different hypotheses about the impact of centra. We show how we create simulation models using a combination of a mathematical model of a fish-like robot using caudal fin propulsion, a propulsion model, and an optimizer, to explore the impact of centra under various scenarios. The optimizer uses the mathematical model to construct valid configurations of the digital robot and uses the utility function and propulsion model to evaluate the performance of each configuration. The evaluations are used to explore the adaptive landscape and find high-performing configurations. Our results show that the high-performing configurations have both increased (flexural) stiffness of the tail and higher tailbeat frequencies.     

The use of environmental DNA of fishes as an efficient method of determining habitat connectivity

This study demonstrated the use of environmental DNA (eDNA) to determine habitat connectivity for migration of fishes between the sea and river. Environmental DNA is DNA fragments released by fishes in water, which can be used as a species-specific marker of the presence/absence of the target species. A year-round water sampling regime at 15 sites on the Yodo River, Japan, was conducted to determine whether three major man-made barriers on the river inhibited the migration of fishes using species-specific detection of DNA fragments from three target migrant species, temperate seabass, Lateolabrax japonicus, flathead grey mullet, Mugil cephalus, and ayu, Plecoglossus altivelis altivelis. The presence/absence of eDNA from target species was consistent with known patterns of species’ seasonal migration. The detection of the DNA of temperate seabass and flathead grey mullet at sites upstream of the dam closest to the river mouth indicated successful upstream migration of these species via a fish ladder bypassing the dam. On the other hand, DNA of these two species was not detected from the upstream side of the two remaining dams, which are not equipped with fish ladders. Ayu is the only species among the three target species with a land-locked population in Lake Biwa located at the headwater of Yodo River. Ayu DNA was detected at most of the sites in the freshwater area during the warm months; however, in the coldest month of February, eDNA was only detected in the uppermost site of Yodo River at the southern tip of Lake Biwa. The eDNA we detected at this site suggests that it was derived from juvenile ayu spending their winter months in the lake. These results suggest that the eDNA analysis presented here can accurately track the seasonal migration of fishes in a river, demonstrating its application as an indicator of habitat connectivity for fishes in association with man-made barriers in a river. The sampling of eDNA involves merely scooping a tank full of water; therefore, it is a simple, rapid, and cost-effective method for long-term monitoring of habitat connectivity associated with the construction of barriers in a river.