Mismatch between fishway operation and timing of fish movements: a risk for cascading effects in partial migration systems

Habitat fragmentation is a growing problem worldwide. Particularly in river systems, numerous dams and weirs hamper the movement of a wide variety of species. With the aim to preserve connectivity for fish, many barriers in river systems are equipped with fishways (also called fish passages or fish ladders). However, few fishways provide full connectivity. Here we hypothesized that restricted seasonal opening times of fishways can importantly reduce their effectiveness by interfering with the timing of fish migration, for both spring‐ and autumn‐spawning species. We empirically tested our hypothesis, and discuss the possible eco‐evolutionary consequences of affected migration timing. We analyzed movements of two salmonid fishes, spring‐spawning European grayling (Thymallus thymallus) and autumn‐spawning brown trout (Salmo trutta), in Norway's two largest river systems. We compared their timing of upstream passage through four fishways collected over 28 years with the timing of fish movements in unfragmented river sections as monitored by radiotelemetry. Confirming our hypothesis, late opening of fishways delayed the migration of European grayling in spring, and early closure of fishways blocked migration for brown trout on their way to spawning locations during late autumn. We show in a theoretical framework how restricted opening times of fishways can induce shifts from migratory to resident behavior in potamodromous partial migration systems, and propose that this can induce density‐dependent effects among fish accumulating in lower regions of rivers. Hence, fragmentation may not only directly affect the migratory individuals in the population, but may also have effects that cascade downstream and alter circumstances for resident fish. Fishway functionality is inadequate if there is a mismatch between natural fish movements and fishway opening times in the same river system, with ecological and possibly evolutionary consequences for fish populations.     

Contemporary ocean warming and freshwater conditions are related to later sea age at maturity in Atlantic salmon spawning in Norwegian rivers

Atlantic salmon populations are reported to be declining throughout its range, raising major management concerns. Variation in adult fish abundance may be due to variation in survival, growth, and timing of life history decisions. Given the complex life history, utilizing highly divergent habitats, the reasons for declines may be multiple and difficult to disentangle. Using recreational angling data of two sea age groups, one‐sea‐winter (1SW) and two‐sea‐winter (2SW) fish originated from the same smolt year class, we show that sea age at maturity of the returns has increased in 59 Norwegian rivers over the cohorts 1991–2005. By means of linear mixed‐effects models we found that the proportion of 1SW fish spawning in Norway has decreased concomitant with the increasing sea surface temperature experienced by the fish in autumn during their first year at sea. Furthermore, the decrease in the proportion of 1SW fish was influenced by freshwater conditions as measured by water discharge during summer months 1 year ahead of seaward migration. These results suggest that part of the variability in age at maturity can be explained by the large‐scale changes occurring in the north‐eastern Atlantic pelagic food web affecting postsmolt growth, and by differences in river conditions influencing presmolt growth rate and later upstream migration.     

The Influence of Discharge on Duration,Ascent Distance,and Fidelity of the Spawning Migration for Paddlefish of the Yellowstone-Sakakawea Stock,Montana and North Dakota,USA

Paddlefish, Polyodon spathula, of the Yellowstone-Sakakawea stock, Missouri and Yellowstone Rivers, Montana and North Dakota, were radio-tagged to assess the influence of spring discharge on duration of river residency, ascent distance, and site-fidelity during spawning migrations of 1999–2002. Contrary to expectations and reported results from other paddlefish populations, fish remained in the river for similar periods of time and ascended to similar reaches in years of higher, more sustained discharge and in years of lower, more fluctuating discharge. In all years, 65 of the 74 migrants (88%) restricted their ascent to reaches below Yellowstone River kilometer (YRkm) 55; only six migrants were found to further ascend to upriver reaches within 20 river kilometers (rkm) of the Intake Diversion Dam (YRkm 114). The lack of detectable annual differences in ascent distance over the study period despite annual differences in Yellowstone River spring flow regimes may have been partially attributed to the apparent site-fidelity demonstrated by the tagged fish over the study period. Ten of the 22 paddlefish contacted in more than one spring migration repeatedly limited their upriver movement to sites that were within 10 rkm of each other. In addition, similar to the reproductive homing tendencies documented in other large-river migratory fishes, site-fidelity occurred in different reaches of the river system. Results from this study suggest that, in years of moderate discharge, site-fidelity may be as influential as the spring flow regime in determining the reaches to which migratory paddlefish ascend. Further research is needed to investigate potential differential spawning success in fish that return to different reaches of the lower Yellowstone River.     

Population dynamics and harvesting of semelparous species with phenotypic and genotypic variability in reproductive age

We study the evolution of polymorphic life histories in anadromous semelparous salmon and the effects of harvesting. We derive dynamic phenotypic and genetic ESS models for describing the evolutionary dynamics. We show in our deterministic analysis that polymorphisms are not possible in a panmictic random mating population. Instead, genetic or behavioral polymorphisms may be observed in populations with assortative mating systems. Positive assortative mating may be supported and generated by behavioral and phenotypic traits like male mate choice, spawning ground selection by phenotype, or within-river homing-migration-distance by size. In the case of an evolutionarily stable dimorphism, the ESS is characterized by a reproductive ideal free distribution such that at an equilibrium the individuals are indifferent from the fitness point of view between the two life histories of early and late reproduction. Different strategy models - that is, phenotypic and genetic ESS models - yield identical behavioral predictions and, consequently, genetics does not seem to play an important role in the present model. An evolutionary response to increased fishing mortality is obvious and may have resource management implications. High sea fishing mortalities drive the populations toward early spawning. Thus it is possible that unselective harvesting at sea may eliminate, depending on the biological system, behavioral polymorphisms or genetic heterozygozity and drive the population to a monomorphic one. If within-river homing migration distances depend on the size of fish, unselective harvesting at sea, or selective harvesting of spawning runs in rivers, may reduce local population sizes on spawning grounds high up rivers. Finally, harvesting in a population may cause a switch in a dominant life-history strategy in a population so that anticipated sustainable yields cannot be realized in practice.     

Spawning migrations in landlocked Atlantic salmon: time series modelling of river discharge and water temperature effects

River discharge and water temperature are frequently cited as controlling the upstream migration of adult salmonids to their spawning areas. The results of earlier studies on the effect of these environmental factors were examined. The statistical methods employed in some of these studies failed to consider the serial correlation often found in migration time series. To assess the effects of discharge and temperature on the migratory activity of the landlocked Atlantic salmon (ouananiche, Salmo salar ), 12 years of data on spawning migrations in the Mistassini R., Quebec, Canada, were analysed and the results obtained by ordinary least squares regression and time series regression were compared. In six of the 12 years, upstream migratory movement was related negatively to changes in river flow, suggesting that fish favour falling water phases for ascent. Water temperature appeared to have little effect on migratory movement. The mean body size of migrating fish decreased significantly throughout the summer; early migrants were on average 11.4% larger (mean fork length 522mm) than late migrants (469 mm). Larger, 3-lake-year salmon migrated 7.2 days earlier than 2-lake-year salmon. Because the residuals from ordinary regression exhibited strong autocorrelation, time series regression was more appropriate than ordinary regression for the analysis of migration time Series.     

Factors influencing the spawning migration of female anadromous brown trout

Radio telemetry was employed to study movements of adult female anadromous brown trout Salmo trutta (sea trout) during upstream spawning migration and following spawning in a stream with tributaries. Sea trout were monitored by manual tracking and by automatic listening stations. The latter suggested that initiation of upstream migration was positively correlated with stream discharge. Individual sea trout performed repeated upstream migration 'initiations'(visits) to areas where they were detected by the automatic listening stations. The first and subsequent upstream migration 'initiations' occurred under conditions of similar water temperature and stream discharge. Manual tracking indicated that in the pre‐spawning state, the distance migrated over 3 days was positively correlated with stream discharge and water temperature, whereas in the post‐spawning state, the total distance migrated was not correlated with any of these two environmental variables.     

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.     

Identifying spawning sites and other critical habitat in lotic systems using eDNA “snapshots”: A case study using the sea lamprey Petromyzon marinus L.

Many aquatic species of conservation concern exist at low densities and are inherently difficult to detect or monitor using conventional methods. However, the introduction of environmental (e)DNA has recently transformed our ability to detect these species and enables effective deployment of limited conservation resources. Identifying areas for breeding, as well as the ecological distribution of species, is vital to the survival or recovery of a conservation species (i.e., areas of critical habitat). In many species, spawning events are associated with a higher relative abundance of DNA released within an aquatic system (i.e., gametes, skin cells etc.), making this the ideal time to monitor these species using eDNA techniques. This study aims to examine whether a “snapshot” eDNA sampling approach (i.e., samples taken at fixed points in chronological time) could reveal areas of critical habitat including spawning sites for our target species Petromyzon marinus. We utilized a species‐specific qPCR assay to monitor spatial and temporal patterns in eDNA concentration within two river catchments in Ireland over three consecutive years. We found that eDNA concentration increased at the onset of observed spawning activity and patterns of concentration increased from downstream to upstream over time, suggesting dispersal into the higher reaches as the spawning season progressed. We found P. marinus to be present upstream of several potential barriers to migration, sometimes in significant numbers. Our results also show that the addition of a lamprey‐specific fish pass at an “impassable” weir, although assisting in ascent, did not have any significant impact on eDNA concentration upstream after the pass had been installed. eDNA concentration was also found to be significantly correlated with both the number of fish and the number of nests encountered. The application of snapshot sampling techniques for species monitoring therefore has substantial potential for the management of low‐density species in fast‐moving aquatic systems.     

White sturgeon (Acipenser transmontanus) passage at the Dalles Dam, Columbia River, USA

White sturgeon (Acipenser transmontanus) ≥95 cm TL were monitored using acoustic and radio telemetry at a large hydroelectric dam (the Dalles Dam) on the Columbia River, during March 2004 through November 2005 to determine timing and routes of passage and to characterize general movements. Transmitters were surgically implanted into 148 fish during the study; 90 were released into the tailrace and 58 into the forebay. We documented 26 passage events by 19 tagged fish: eight upstream via fish ladders and 18 downstream, mostly through open spill gates. During the study 17 fish entered the two ladders one or more times; 11 entered only the east ladder, three entered only the north ladder, and three entered both ladders at sometime. Residence time within the ladders by individual fish was variable, ranging from about 1 min to nearly 6 months (median = 7.7 h). Only six fish successfully ascended the east ladder, one fish twice. We could not unequivocally determine which fish ladder one fish used to pass upstream. Differences in construction between the north and east fish ladders may account for the greater success of the east fish ladder in passing sturgeon upstream. Changes to operations at hydroelectric dams to benefit migrating anadromous salmonids may influence upstream or downstream passage by white sturgeon. Altering patterns and timing of spill discharge, altering fish ladder entrance attraction flows, and the use of lights, sound, and partial barriers to direct other species of fish to preferred passage routes have unknown effects on sturgeon passage. A better understanding of the consequences to the metapopulation of increasing or precluding upstream or downstream passage is needed.     

Swimming performance and morphology of juvenile sockeye salmon, <Emphasis Type="Italic">Oncorhynchus nerka</Emphasis>: comparison of inlet and outlet fry populations

We raised two populations of sockeye salmon fry from fertilized eggs in the laboratory and tested the hypothesis that outlet fry populations, fish which must migrate upstream to reach rearing lakes after yolk-sac absorption, have better swimming ability and morphological characteristics conducive to enhanced swimming performance than inlet fry populations, fish which migrate downstream to rearing lakes. Despite being of identical age, fry from the outlet population were larger (approx. 6.7% longer, ~5 mm on average) and more laterally compressed than inlet fry at the time of our initial experiments. Using an open-top box flume, we found that the burst-swimming performance (in cm s⁻¹) of the outlet population was 31% better. We found no differences between populations in prolonged-swimming performance. We were unable to find any direct relationships between measures of swimming performance and size or shape variables, suggesting that the larger, more robust morphology of outlet fry was not responsible for the superior burst ability. Recent biochemical studies indicate outlet fry may be metabolically better provisioned for burst swimming than inlet fry. It is possible that the morphological differences between the populations of fry reflect adaptations needed by adults during their migration and spawning.     

Assessing the relative importance of temperature,discharge, and day length on the reproduction of an anadromous fish (Alosa alosa)

1. Climate change threatens anadromous fishes such as the allis shad (Alosa alosa) populations of which have declined since the 20th century in Europe. Sensitivity to climate change could be quantified by determining the fish’s spawning behaviour, defined as the timing of reproduction (i.e. spawning events) as a function of temporally variable environmental factors. The cues that fish use to time reproduction could determine their response to climate change.
2. A machine learning technique (boosted regression tree) was calibrated using a 14-year dataset composed of daily measures of environmental factors and fish occurrences during reproduction. The boosted regression tree provides complete insight into the complex relationship between the spawning probability, i.e. the probability for a fish to reproduce, and environmental factors that might evolve with climate change.
3. The spawning probability was positively related to day length (44.6%) and water temperature (34.7%) and negatively related to river discharge (20.7%). Optimal reproductive conditions corresponded to a difference in day length between 0 and 0.04 hr, a water temperature between 15 and 26°C and a river discharge between 55 and 665 m³/s; these conditions are currently being utilised by allis shad populations in the Garonne and Dordogne rivers, France.
4. This study highlights the relative influence of environmental factors on the observed spawning period as well as the evolution of habitat suitability during the 14-year period. The novelty of this study stems from assessing population process data, i.e. the occurrence of fish reproduction, rather than mere occurrence data in an ecological niche model study. Climate change may lead to a shift in spawning phenology, as the water temperature and river discharge will also change. Therefore, conservation plans need to integrate these effects on spawning grounds.

     

Hydrological Status of the spawning ground of Acipenser sinensis underneath the Gezhouba Dam and its relationship with the spawning runs

The spawning runs of Chinese Sturgeon (CS; Acipenser sinensis) were observed 37 times below Gezhouba Dam of Yangtze River between 1983 and 2004. Five hydrological factors (water temperature, water level, flow discharge, silt content and current velocity) were monitored on a daily basis at the spawning ground between October and November for 22 consecutive years (1983–2004). The effect of current velocity on the spawning ground at the bottom layer of the river, where CS was spawning for four years, was measured (1996–1999). The authors of this study analyzed the relationship between the five hydrological factors and the respective spawning runs. Twenty-two years of continuous observations indicated that the daily mean values of all the five hydrological factors fluctuated within a certain range when CS was spawning. It was concluded that the optimal values for the hydrological factors during the spawning runs are 18.0–20.0°C for temperature, 14100 m³/s for discharge volume, 42.0–45.0 m for water level above the sea level, and 0.2–0.3 kg/m³ for silt content in the water, wherein the current velocity above the bottom layer to stimulate the fish to spawn should be between 1.0–1.7 m/s. The optimal water temperature might provide an essential precondition for other factors to trigger spawning. As water temperature reaches the optimal values and most of the other parameters are at the brink of deviation from their optimal range of values (water level, current velocity and silt content in the water), CS would begin to spawn. By 2009, when the Yangtze Three Gorges Project, which is located 45 km upstream of the Gezhouba Dam, is completed and begins to operate normally, changes in the downstream water temperature are expected to occur, which may have a negative effect on the development of gonad and the stimulation of spawning of CS; however, the anticipated decrease of the silt content in the water may be considered favorable for the performance of the spawning site.     

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.     

Migrations and swimming capabilities of endangered pallid sturgeon (Scaphirhynchus albus) to guide passage designs in the fragmented Yellowstone River

Fragmentation of the Yellowstone River is hypothesized to preclude recruitment of endangered Scaphirhynchus albus (pallid sturgeon) by impeding upstream spawning migrations and access to upstream spawning areas, thereby limiting the length of free‐flowing river required for survival of early life stages. Building on this hypothesis, the reach of the Yellowstone River affected by Intake Diversion Dam (IDD) is targeted for modification. Structures including a rock ramp and by‐pass channel have been proposed as restoration alternatives to facilitate passage. Limited information on migrations and swimming capabilities of pallid sturgeon is available to guide engineering design specifications for the proposed structures. Migration behavior, pathways (channel routes used during migrations), and swimming capabilities of free‐ranging wild adult pallid sturgeon were examined using radiotelemetry, and complemented with hydraulic data obtained along the migration pathways. Migrations of 12–26% of the telemetered pallid sturgeon population persisted to IDD, but upstream passage over the dam was not detected. Observed migration pathways occurred primarily through main channel habitats; however, migrations through side channels up to 3.9 km in length were documented. The majority of pallid sturgeon used depths of 2.2–3.4 m and mean water velocities of 0.89–1.83 m/s while migrating. Results provide inferences on depths, velocities, and habitat heterogeneity of reaches successfully negotiated by pallid sturgeon that may be used to guide designs for structures facilitating passage at IDD. Passage will provide connectivity to potential upstream spawning areas on the Yellowstone River, thereby increasing the likelihood of recruitment for this endangered species.     

Poor water quality constrains the distribution and movements of twaite shad <Emphasis Type="Italic">Alosa fallax</Emphasis><Emphasis Type="Italic">fallax</Emphasis> (Lacépède, 1803) in the watershed of river Scheldt

Worldwide, river fragmentation is primarily responsible for the decline of populations of migrating fish. In particular, anadromous fish species, which necessarily migrate to fresh water to reproduce, are endangered since many are no longer able to reach their natural spawning sites. In addition, pollution of rivers effectively prevents upstream or downstream movements and blocks access to spawning grounds. This article investigates how poor water quality interferes with the life history cycle of twaite shad Alosa fallax fallax (Lacépède, 1803), an anadromous clupeid fish, in the watershed of River Scheldt, a heavily impacted environment in West Europe. We used two models based on known ecological and environmental information to explain past and present twaite shad distribution within the watershed and to make inferences about a future population recovery and juvenile habitat value. We demonstrated that historical spawning areas satisfy water quality conditions necessary to support spawning and successful development of early life history stages of the twaite shad. However, poor water quality conditions just upstream the freshwater–saltwater boundary still act as an effective migration barrier for upstream movement. As a consequence, spawning grounds are inaccessible and the population is dominated by seasonal adults occurring in the lower estuarine part of the watershed. This article provides testable and diagnostic information to the watershed management in that it identifies habitat and water quality requirements needed to support the expected recovery of an endangered anadromous fish population. Guest editors: S. Dufour, E. Prévost, E. Rochard & P. Williot Fish and diadromy in Europe (ecology, management, conservation)     

NATURAL REPRODUCTION AND HYDROLOGIC REQUIREMENTS OF PELAGOPHIL FISH IN THE THREE GORGES RESERVOIR北大核心CSCD

To reveal the relationships between spawning activities of pelagophil fish and environmental factors in the Three Gorges Reservoir (TGR), fish eggs and larvae were sampled at Zhongxian section, from May to July in 2016 and 2017. 60 fish species including 30 pelagophil fishes were sampled. Hemiculter bleekeri and Squalidus argentatus were the dominant species in eggs, and Hemiculter and Pseudolaubuca were abundant in larvae. In 2016, 7.041 billion eggs and 103.763 billion larvae drifted through the sampling section, and in 2017, 3.454 billion eggs and 34.427 billion larvae drifted through the sampling section. The amount of eggs and larvae of the four major Chinese carps reached 2.909 billion in 2016 and 2.117 billionin in 2017. Spawning activities of pelagophils fish varies temporally in different river reaches, which mainly occurred after early-June in the river reach from Zhongxian to Fuling but begun in mid-May in the river reach upstream of Changshou. Fish eggs and larvae from the river reach upstream of Changshou provided a large resource for recruitment to the fishery in TGR. Classification and regression trees (CART) analysis explored that spawning activities of the four major Chinese carps were favord when water discharge was rasing. But, spawning activities of Hemiculter bleekeri and Squalidus argentatus had no relationship with water discharge. In order to protect the spawning grounds and meet environmental requirements for natural reproduction of pelagophils fish, we propose ecological regulation of the cascade hydro-power projects in the Jinsha River and maintenance of large free-flowing stretches upstream of TGR. © 2019, Institute of Hydrobiology, Chinese Academy of Sciences. All rights reserved.     

Migratory and reproductive behaviour of female adult Atlantic salmon, Salmo salar L., in a spawning stream

Migration and spawning behaviour of eight Atlantic salmon adult females were analysed by radio-tracking in relation to the degree of sexual maturity in a spawning tributary of the R. Sélune. Six of them were grilse and four of them were ripe. All the fish migrated upstream until reaching their spawning site at a distance of 4–12 km from the trap. The daily migration rate up to this site was inversely correlated with the length of the female. Spawning occurred in all fish at the same time when the water temperature increased dramatically. Spawning lasted 1–10 days according to the fish. After spawning, females quickly moved downstream only small distances and then stayed in approximately the same location until death. This study concluded that ripeness did not influence behaviour on the spawning migration and describes certain characteristics of the reproductive phase in a spawning tributary.     

Availability of and access to critical habitats in regulated rivers: effects of low-head barriers on threatened lampreys

1.  Conservation of freshwater animal populations requires their access to, as well as sufficient availability of, critical habitats, such as those for reproduction. Abundant small-scale barriers may cause extensive fragmentation of freshwater habitat but, by comparison to larger structures their effects are rarely considered by catchment managers. The relationship between the distribution of, and access to, spawning habitat in a regulated river, characterized by abundant small barriers, was examined for river lamprey Lampetra fluviatilis , a threatened migratory fish.
2.  Telemetry of adult lamprey in the River Derwent, North East England was used to quantify upriver migration and access to spawning habitat, together with surveys of spawning habitat availability and spawning activity between 2002 and 2007.
3.  Access in to the Derwent appeared severely restricted by a tidal barrage, beyond which lamprey migrated rapidly in unobstructed reaches. Of all lamprey tagged in the lower 4 km of river, or ascending the barrage, 64% and 17% passed the first and second weirs respectively, with high flows crucial for this. Although over 98% of lamprey spawning habitat occurred more than 51 km upstream, on average just 1.8% of river lamprey spawners were recorded there.
4.  In order to protect or rehabilitate species or species assemblages, greater attention needs to be paid to the relative spatial distribution of low-head barriers and the resultant availability of key habitats within individual catchments. This is particularly important given the renewed emphasis internationally on low-head hydropower solutions as a source of renewable energy, and the rapid growth in numbers of low-head barriers in many catchments.     

Seasonal movement and distribution of fluvial adult bull trout in selected watersheds in the mid-Columbia River and Snake River basins

From 1997 to 2004, we used radio telemetry to investigate movement and distribution patterns of 206 adult fluvial bull trout (mean, 449 mm FL) from watersheds representing a wide range of habitat conditions in northeastern Oregon and southwestern Washington, a region for which there was little previous information about this species. Migrations between spawning and wintering locations were longest for fish from the Imnaha River (median, 89 km) and three Grande Ronde River tributaries, the Wenaha (56 km) and Lostine (41 km) rivers and Lookingglass Creek (47 km). Shorter migrations were observed in the John Day (8 km), Walla Walla (20 km) and Umatilla river (22 km) systems, where relatively extensive human alterations of the riverscape have been reported. From November through May, fish displayed station-keeping behavior within a narrow range (basin medians, 0.5-6.2 km). Prespawning migrations began after snowmelt-driven peak discharge and coincided with declining flows. Most postspawning migrations began by late September. Migration rates of individuals ranged from 0.1 to 10.7 km/day. Adults migrated to spawning grounds in consecutive years and displayed strong fidelity to previous spawning areas and winter locations. In the Grande Ronde River basin, most fish displayed an unusual fluvial pattern: After exiting the spawning tributary and entering a main stem river, individuals moved upstream to wintering habitat, often a substantial distance (maximum, 49 km). Our work provides additional evidence of a strong migratory capacity in fluvial bull trout, but the short migrations we observed suggest adult fluvial migration may be restricted in basins with substantial anthropogenic habitat alteration. More research into bull trout ecology in large river habitats is needed to improve our understanding of how adults establish migration patterns, what factors influence adult spatial distribution in winter, and how managers can protect and enhance fluvial populations.     

RHEOTROPISM IN FISHES

(1) The fluid properties of air and water enable animals to orientate to flow and this behaviour in water is termed rheotaxis. Fish, however, have a wide range of responses to currents, extending beyond simple orientation, and the term rheotropism is therefore used as a ‘portmanteau’ word to describe all such reactions. (2) Fish detect currents directly by flow over the body surface or indirectly by other stimuli. Indirect responses are more common and occur in response to visual, tactile and inertial stimuli resulting from displacement of the fish by the current. Reactions to displacement of visual images are called optomotor reactions. The lateral line is not involved except in the detection of small localized jets of water. It has not been demonstrated that any fish can detect the current by electrical stimuli, although it is theoretically possible for some to do so. (3) In the basic form of rhotaxis the fish heads upstream and maintains station by stemming the current. Current detection thresholds fall within the range 0.4 to 10 cm/s for tactile stimuli but may be as low as 0.03 cm/s for visual stimuli. (4) Visual responses have been studied by simulating displacement by the current in optomotor apparatus. Fish respond to a rotating black-and-white-striped background by compensatory movements of the head and eyes - optokinetic nystagmus - or by the optomotor reaction, in which the fish swims with the background. (5) Fish show an orthokinesis in optomotor apparatus, their mean swimming speed increasing with the speed of rotation of the background. The precise form of the relationship varies between species and there is also considerable individual variation in performance. Fish accelerate and decelerate relative to the background, fixating on a particular stripe for short periods. (6) Factors limiting the appearance of the optomotor response are contrast, illuminance, acuity, critical flicker fusion frequency and spectral sensitivity. (7) Fish tolerate retinal image movements equivalent to those received when they are carried forwards by the current but not to those received when they are carried backwards. There are ganglion cells in the optic tectum which are sensitive to the direction of movement of targets across the visual field. In the goldfish there are significantly more units sensitive to movements in the temporo-nasal than in the opposite direction. (8) There are close parallels between the behaviour of fish in schools and in an optomotor apparatus. The optomotor response is apparently innate, occurring in newly hatched fry. (9) Physical and chemical factors can modify rheotaxis. Temperature and olfactory stimuli affect both the sign of the taxis and the kinetic component of the behaviour. (10) Thyroid hormones which are involved in the control of migration have been shown to affect the kinetic component of rheotaxis. (11) Fish show a number of hydrodynamic adaptations to life in currents. Morphological modifications are greatest in fish from torrential streams, which show extreme dorsoventral flattening and have specialized adhesive organs. Other fish select areas of low velocity or decrease their buoyancy with increasing current speed. (12) Rheotropic behaviour plays an important role in the distribution of fish within stream systems, in the maintenance of territory and station and in feeding behaviour. Territory, station and spawning sites in salmonids are all selected in relation to water velocity. (13) Water currents are thought to provide either a transport system or directional clues for fish on migration. The fish either does not respond to the current and is carried passively downstream, or it makes an orientated movement, swimming up- or downstream. (14) Eggs and larvae are known to drift passively downstream from their spawning grounds and some adult fish may also drift passively. In the sea both adult and juvenile fish use a form of modulated drift associated with vertical migration. Fish move up into midwater either by direct tidal selection or in relation to the diel cycle of illuminance. In fresh water the downstream migrations of salmonid fry, and smolts under some conditions, occur by modulated drift. (15) There is no evidence that fish migrating in the sea orientate to the current, but in fresh water the upstream migrations of diadromous fish are clearly orientated movements. (16) Water velocity is a major factor for salmonids migrating upstream. For fry it limits the occurrence of upstream migrations and for adults it can also prevent upstream movement. But migrations are often initiated by freshets, and changing water velocity is thought to be the most important factor associated with a freshet. (17) Both environmental and genetic factors affect the direction of migration in relation to the current. In some sockeye salmon fry direction is determined by temperature, but in others the overall direction of movement is genetically determined and environmental factors only modify the behaviour. (18) Rheotropic behaviour has a number of important practical applications in the capture of fish and in guiding them past dams and power stations. (19) The optomotor response plays a basic role in the capture of roundfish by trawls under conditions when the fish can see the gear. Many fish are caught because they become fatigued after a prolonged period of swimming at the same speed as the trawl. (20) Most success in guiding fish away from hazardous areas and bypassing them round dams has been achieved with mechanical barriers which depend on rheotropic reactions of the fish. (21) Louvre screens are very successful in deflecting juvenile salmonids migrating downstream past small dams but are impracticable at large dams. Instead, the turbine intakes are commonly sited at a considerable depth and fish are bypassed by mechanical screens either at the surface of the forebay or into the gatewells immediately upstream of the turbine intakes. (22) With upstream migrants the basic problem is to attract fish to the lower end of the fishways. An adequate ‘attraction velocity’ is an important feature of fishways, which must be sited so that the fish avoid the high velocity discharges from spillways and turbines.