Potential negative impacts of introduced rainbow trout on endangered Sakhalin taimen through redd disturbance in an agricultural stream, eastern Hokkaido

Nomoto K, Omiya H, Sugimoto T, Akiba K, Edo K, Higashi S. Potential negative impacts of introduced rainbow trout on endangered Sakhalin taimen through redd disturbance in an agricultural stream, eastern Hokkaido.
Ecology of Freshwater Fish 2010: 19: 116–126. © 2009 John Wiley & Sons A/S
Abstract –  Sakhalin taimen ( Hucho perryi ) populations have decreased in Hokkaido, northernmost Japan, primarily because of overexploitation and habitat degradation. Here we document another threat to this species, that of spawning redd superimposition by artificially introduced rainbow trout ( Oncorhynchus mykiss ). Sakhalin taimen and rainbow trout are the only spring-spawning salmonid species in Hokkaido. In 2006–2008, spawning activities of both these species were observed in a Hokkaido stream, and it was determined that their spawning periods overlapped during mid-late April. They also spawned at similar water velocities, depths and substrate compositions. Although female Sakhalin taimen were larger than female rainbow trout, their egg burial depths were nearly identical. During the observation period, rainbow trout redds were approximately five times more abundant than Sakhalin taimen redds, and about 30% of the observed Sakhalin taimen redds were superimposed by rainbow trout redds. The high degree of spatial and temporal overlap in spawning, the similar egg burial depths of both species, and the high proportion of superimposed redds suggest that the introduced rainbow trout impact the endangered Sakhalin taimen in Hokkaido, and possibly, in other areas where the two species occur together.     

Brown trout redd superimposition in relation to spawning habitat availability

Abstract – The relationship between redd superimposition and spawning habitat availability was investigated in the brown trout (Salmo trutta L.) population inhabiting the river Castril (Granada, Spain). Redd surveys were conducted in 24 river sections to estimate the rate of redd superimposition. Used and available microhabitat was evaluated to compute the suitable spawning habitat (SSH) for brown trout. After analysing the microhabitat characteristics positively selected by females, SSH was defined as an area that met all the following five requirements: water depth between 10 and 50 cm, mean water velocity between 30 and 60 cm s^?1, bottom water velocity between 15 and 60 cm s^?1, substrate size between 4 and 30 mm and no embeddedness. Simple regression analyses showed that redd superimposition was not correlated with redd numbers, SSH or redd density. A simulation‐based analysis was performed to estimate the superimposition rate if redds were randomly placed inside the SSH. This analysis revealed that the observed superimposition rate was higher than expected in 23 of 24 instances, this difference being significant (P < 0.05) in eight instances and right at the limit of statistical significance (P = 0.05) in another eight instances. Redd superimposition was high in sections with high redd density. High superimposition however was not exclusive to sections with high redd density and was found in moderate‐ and low‐redd‐density sections. This suggests that factors other than habitat availability are also responsible for redd superimposition. We argue that female preference for spawning over previously excavated redds may be the most likely explanation for high superimposition at lower densities.     

Competitive exclusion of brown trout Salmo trutta L., by rainbow trout Oncorhynchus mykiss Walbaum, in lake tributaries, New Zealand

Spawning runs of brown trout Salmo trutta L., and rainbow trout, Oncorhynchus mykiss Walbaum, were compared in three tributaries of two lakes for timing, numbers and redd characteristics. Redd characteristics for the two species in four other streams were examined. No significant differences in redd characteristics between the two species were observed, but in all lake tributaries the possibilities of superimposition of spring spawning rainbow trout on redds of brown trout were examined. A common feature was the high ratio of numbers of spawners to the area of preferred spawning gravel, and it was concluded that in the lake tributaries, superimposition resulted in increased mortality of brown trout eggs, and in one case elimination of the brown trout population. Management possibilities for control of the species proportions in comparable situations are discussed.     

A fish selective obstacle to prevent dispersion of an unwanted fish species, based on leaping capabilities

Abstract  The European minnow, Phoxinus phoxinus (L.), constitutes a serious threat to natural brown trout, Salmo trutta L., stocks in Norway following its introduction and translocation. In the present study, the leaping capabilities of the European minnow (50–110 mm total length) and brown trout (64–255 mm total length) were investigated with the aim of constructing suitable waterfall barriers to prevent further unwanted dispersal of the minnow, but still allow passage of larger brown trout. No successful leap of minnows was recorded at 4.9–6.5 °C, even at height intervals as low as 3 cm. At 14.0–16.5 °C, minnows were able to force waterfall barriers up to 27 cm high. At 4.9–6.5 °C brown trout forced waterfall barriers up to 40 cm. Thus, building of such barriers in brooks and rivers can help prevent the dispersal of minnows, and still allow larger brown trout to pass during feeding and spawning migrations.     

Spawning period of a southern brown trout population in a highly unpredictable stream

Abstract –  The spawning period of brown trout ( Salmo trutta L.) was studied in the river Castril, southern Spain, by means of redd counts. This mountain stream is located near to the southern limit of the species' natural distribution range and it shows a highly unpredictable flow regime. The spawning period extended from December to mid-April and the maximum reproductive activity was in February. These results represent the latest reproduction date and the longest spawning period reported in the literature along the natural distribution range of the species. However, belated spawning in the Castril is congruent with the known latitudinal cline: the lower the latitude, the later the spawning period. Our results, along with a review of the literature on natural populations, also showed that the duration of reproduction is the longer, the lower the latitude. Spawning lasted twice as long in the main stem of the river, which is connected with a reservoir, than in the isolated reaches. These differences may be linked to the influence of the reservoir and to habitat fragmentation. We discuss and support the hypothesis that a long spawning period is an advantage for survival in unpredictable habitats. The belated and protracted spawning period found in river Castril has important implications in fisheries management. A strong research effort is needed in order to fill the critical lack of data on southern brown trout populations.     

Thermal regime in the hyporheic zone of two contrasting salmonid spawning streams: ecological and hydrological implications

The temperatures of stream water and hyporheic water were monitored in spawning areas of an upland stream (Girnock Burn) and a degraded lowland stream (Newmills Burn). Both streams are located in North-east Scotland and are spawning streams for Atlantic salmon, Salmo salar L., and brown trout, S. trutta L. Monitoring between the spawning and hatching of ova revealed significant differences in the thermal regimes of the two sites. Whilst temperatures in stream and hyporheic water were similar in the Girnock, notable attenuation in the Newmills Burn resulted in increasing moderation of thermal regime with hyporheic depth. This probably reflects contrasting groundwater–surface water interactions in the two systems. In the Girnock an open gravel stream bed and lack of groundwater influence resulted in stream water downwelling through the gravel. In contrast, fine bed sediments and local groundwater inputs resulted in more complex stratification of temperatures in Newmills Burn. However, even in the conditions of the Newmills Burn, temperature based salmonid embryo development models predicted that egg burial depth has an insubstantial effect on ova development.     

Male antagonistic behaviour after spawning suggests paternal care in brown trout,Salmo trutta

Abstract – The evolution of parental care depends on the sex‐dependent resolution of a trade‐off between present and future reproduction. In salmonids, the mating systems are often characterised by fierce male–male competition, and the absence of paternal care is widely admitted in these species. However, the fact that brown trout eggs can be cannibalised by peripheral individuals just after spawning suggests that dominant males would benefit from protecting their offspring by keeping cannibals away from the nest. We used 77 field observations of brown trout spawning to document the defensive behaviour of dominant males towards peripheral individuals before and after spawning, and test whether this behaviour was related to the probability of the clutch being cannibalised. We showed that dominant males stayed on the nest and chased peripherals even after the eggs were laid and fertilised. The number of chases performed by dominant males during the 2 min following spawning was negatively correlated to the probability of egg cannibalism. This result indicates that brown trout males can provide direct benefits to females through protection against egg cannibalism by peripheral individuals.     

Water level influences migratory patterns of anadromous brown trout in small streams

Habitats modify the population ecology of species. Here, we show how low water level influences abundance and size of adult anadromous brown trout (Salmo trutta) entering a small, South Norwegian stream for spawning. After smolting, the fish appear chiefly to feed within 10 km of the home stream. In the autumn, South Norwegian streams typically flood because of heavy rainfall, when the anadromous brown trout entered from the sea. Mean annual duration of the upstream migration period was 34 days and ended when the flooding ended and the water temperature dropped to below 4°C. During most of the migration period, on average two trout ascended the river per day. The sexes entered the spawning area concurrently, and the male:female ratio of the anadromous trout was 1.27. No fish entered when the water depth just downstream of the spawning area was below 5 cm, and mean number of fish increased with increasing water depth to ca. 30 cm, but not at higher flows when the ascent gradually decreased. Mean and maximum size of the entering spawners increased with water depth between 5 and 16 cm. Among those that had been to sea, most were recaptured in the home stream, 4% in other streams, but only two of the strays were caught close to spawning time. The present results illustrate that population traits of anadromous brown trout from a small stream differ from those in larger rivers, probably because of selection associated with water flow.     

Factors influencing the success of salmonid egg development in river substratum

Interstitial water conditions in the hyporheic zone of the stream bed are determinants of salmonid egg hatching success. We used standardised egg exposures to develop and validate discriminant analysis and generalised linear model models linking the hatching success of brown trout (Salmo trutta) with physicochemical factors of the interstitial zone (e.g., oxygen, specific conductance, nitrate, nitrite, ammonium, pH and redox potential). Interstitial water quality was identified as a limiting factor for egg development (median of relative hatching rates = 0.23). Hatching success was unimodal in hatchery and field references incubated in free‐flowing water, but bimodal (very high or very low hatching success) in natural sediment exposures. The effects of physicochemical factors on the hatching success of Salmo trutta strongly depended on both the time and spatial scale analysed. The variables retained in the models differed between the macroscale (over all rivers), the river‐specific scale (within a river) and the microscale (at different sediment depths). Egg hatching success decreased with increased substratum depth (decrease of 26% in 150 mm compared with 50 mm). Increasingly more variable interstitial water conditions (e.g., oxygen) throughout the egg incubation period suggest progressive degradation rates in the stream substratum during the incubation period at the microscale level. Consequently, consideration of different spatial and temporal scales is necessary for the evaluation of habitat quality in salmonid conservation and catchment management plans.     

Behaviour and growth of juvenile brown trout exposed to fluctuating flow

Abstract –  Parr of brown trout ( Salmo trutta L.) were exposed to constant or fluctuating water level and flow in artificial outdoor stream channels during 21 days in three consecutive runs, and their individual behaviour and social interactions were monitored. Despite the large variation in flow, no clear differences in growth, which was generally poor, and behaviour were found between treatments. The signs of a social structure in the stream channels were very subtle and there were no clear relation between growth rate and aggressive/dominant behaviour in any of the treatments in accordance with findings in studies of growth and dominance in the wild. Thus, in regulated rivers, daily fluctuations in flow and water level (hydropeaking) are not likely to alter the behaviour of juvenile brown trout or affect their energy turnover negatively. Several studies now indicate that if stranding can be avoided, hydropeaking has relatively small direct effects on stream salmonids. Therefore, future research should focus on cascading ecological effects that may influence the productivity of the ecosystems in regulated rivers.     

Using underwater cameras to describe the reproductive behaviour of the endangered eastern freshwater cod Maccullochella ikei

Abstract –  Underwater cameras were used to observe the breeding behaviour of the endangered eastern freshwater cod, Maccullochella ikei , over 3 years and across three areas in the Mann and Nymboida rivers, Australia. The annual breeding season for M. ikei was short and succinct, lasting only 8–10 weeks. Spawning commenced each year in the lowest altitude area during the first week of spring, and approximately 1 week later in the closest upstream area. Day-length is the primary spawning cue for M. ikei , but increasing water temperature may also be of importance. Nesting sites were located in slow-flowing pools, under cover such as large boulders and bedrock shelves, at depths of 0.9–4.0 m, and with one or two entrances only. The nesting site was vigorously cleaned by the male up to 1 week prior to spawning and was only entered by the female for spawning. Paternal care of eggs and larvae was undertaken for up to 24 days, after which larvae dispersed. Greater protection of breeding M. ikei must be a management priority to ensure long-term conservation of the species.     

The influence of hydrology on the recruitment of brown trout in an Alpine river,the Ybbs River,Austria

Abstract – We monitored yearly recruitment (1997–2008) of brown trout (Salmo trutta) in a fourth‐order Austrian Alpine river. The relative proportion of recruits to adult fish varied strongly among years (5.6–66.4%). These proportions were strongly correlated with specific flow patterns. High flows before and during the spawning period were positively correlated with recruitment, whereas high flows during incubation and emergence were negatively correlated with recruitment success. Unsteady flow modelling supported a causal hypothesis for these relationships in demonstrating that discharges > 30 m³·s^?1 resulted in substantial sediment motion (erosion and deposition) in suitable spawning areas within the study stretch.     

Migration of landlocked brown trout in two Scandinavian streams as revealed from trap data

Abstract –  Anthropogenic barriers that may interfere or prevent fish migration are commonly found in streams throughout the distribution of salmonids. Construction of fish passages in streams is a common solution to this problem. However, the goal with fish passages is often, at least in Scandinavia, to allow Atlantic salmon ( Salmo salar L.) and migratory brown trout ( S. trutta L.) to get access to spawning areas above these barriers. Hence, the fish passages may often only be open during the spawning migration of salmonids (late summer to autumn). We present data, on wild brown trout migration, from two trapping systems in two Scandinavian streams showing that intra- and interstream migrations are common throughout the summer and autumn. Moreover, differences in size were found between trap-caught trout and electrofished trout where trapped trout were generally larger than electrofished trout. We suggest that the current regime with fish passages only open parts of the year can have negative effects on populations by depriving trout from the possibility to perform migrations throughout the year.     

Micro‐scale distribution of brown trout: an opportunity for kin selection?

Abstract – Fourteen microsatellite loci were used to analyse the degree of relatedness between and within cohorts of stream-living brown trout ( Salmo trutta L.). Differences in degree of relatedness were found between cohorts with mature and older having the highest intracohort relatedness. The higher genetic relatedness among the mature and older trout is interpreted as indications of clustering of related mature and older trout prior to spawning in the vicinity of spawning grounds. Young-of-the-year trout showed a lower degree of association with related trout of the same cohort than did mature and older trout. It is argued that the clustering of mature and older trout may facilitate kin selection and that the low degree of association among related young-of-the-year trout could be explained by juvenile fish avoiding competition for territories with related individuals.     

The influence of water depths and inter-specific interactions on cover responses of juvenile Atlantic salmon

Abstract  – The responses of salmon parr, Salmo salar, to instream cover, related to several water depths, were tested in an ellipsoidal stream tank. Opaque plastic covers, most of which were 20 cm in length and 16 cm high, were randomly distributed through the tank, occupying about 3% of the bottom area. Six salmon parr were used for each experiment. In experiments with channel depths of 40 cm, the proportion of salmon under covers was 36.4%, at 30 cm 60.7%, and at 20 cm, 79.4%. Presence of similar-sized brown trout, Salmo trutta, increased the use of covers in channels and greater use of the pool area by salmon. The salmon were completely dominated by the trout, making about four times more more aggressive than salmon. Our findings show that water depth itself provides cover for young salmon.     

Differences in longitudinal distribution patterns along a Honshu stream of brown trout <Emphasis Type="Italic">Salmo trutta</Emphasis>, white-spotted charr <Emphasis Type="Italic">Salvelinus leucomaenis</Emphasis> and masu salmon <Emphasis Type="Italic">Oncorhynchus masou</Emphasis>

Brown trout Salmo trutta were first introduced into Japan in 1892, and they currently naturally reproduce in several rivers in Honshu and Hokkaido, Japan. Although negative impacts of brown trout introductions on native salmonid fishes have been documented in some Hokkaido rivers, studies of ecological interactions between brown trout and native salmonid fishes on Honshu are limited. In this study, we describe the longitudinal distribution patterns of introduced brown trout, white-spotted charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou in a 4 km stretch of a stream in central Honshu. Underwater observations were conducted in all pools within upstream, middle and downstream sections (190–400 m in length) of this stretch in order to estimate the densities of these species. Only white-spotted charr was observed in the upstream section, while brown trout and masu salmon were observed in the middle and downstream sections. Masu salmon densities, however, were much lower than brown trout densities. In the downstream section, white-spotted charr was absent. These results are consistent with results from previous studies of Hokkaido rivers, where it was found that white-spotted charr in low-gradient areas tend to be displaced by brown trout.     

High variability in spawning migration of sea trout,Salmo trutta,in two northern Swedish rivers

Abstract Telemetry was used to examine spawning migration of sea trout, Salmo trutta L. (n = 126), in two rivers in northern Sweden. The spawning areas defined by radio‐tagged fish differed between the river systems. In the River Vindelälven, sea trout spawned in the main stem and 80% of tagged individuals returned to areas where hatchery‐reared juveniles had been previously stocked. In the River Piteälven, 74% of tagged sea trout ascended tributaries for spawning. Tagged fish were categorised into three groups of migratory pattern. cart (classification and regression tree) analysis indicated that distance from tagging location to spawning site (S_dist) explained the migratory patterns. Large S_dist separated fish with stepwise upstream migration from those with up‐ and downstream migrations and one‐directional direct migration. Fish tagged early in the season migrated the longest distance to spawning areas. Stocking locations and sex explained the large search behaviour up‐ and downstream in the rivers. The findings are important for the sustainable management of sea trout in the Gulf of Bothnia.     

Genetic differentiation between anadromous and freshwater resident brown trout (Salmo trutta L.): insights obtained from stable isotope analysis

Abstract –  A genetic survey was carried out on recently emerging fry to investigate interbreeding between anadromous and freshwater resident brown trout in La Roche Brook, a major spawning tributary of the Oir River system (Normandy, France). Emerging fry were sampled at three different sites in spring 2002 and 2003. Stable isotope analysis (SIA) allowed the classification of each individual as the progeny of resident ( N  = 76) or anadromous ( N  = 58) female trout. Microsatellite DNA analysis showed that genetic diversity between fry samples can be explained by microgeographical (various sampling sites) and temporal (2 years of sampling) variations rather than maternal origin (anadromous vs. freshwater resident). We conclude that a high level of gene flow exists between the two morphs when anadromous adults have access to the spawning grounds of residents. We highlight the relevance of using SIA in conjunction with genetic analysis to study interactions between the two morphs.     

Stability in reproductive timing and habitat usage of Chinook salmon across six years of varying environmental conditions and abundance

The spatial and temporal distribution of spawning activity by autumn‐run Chinook salmon Oncorhynchus tshawytscha (Walbaum) was examined across multiple years. The study period included two years of extreme drought conditions when water temperatures in the spawning reach of the Stanislaus River were considered sub‐optimal for spawning and egg incubation. Despite varying levels of superimposition, redd counts and densities remained stable and positively associated with river location, indicating that superimposition may be driven by habitat preference rather than the absence of suitable spawning locations. Spawning occurred slightly later during drought years (6–10 days later compared with wetter years). This delay was attributable to deferred migration instead of deferred spawning, and the majority of redds were constructed at water temperatures exceeding the optimal temperature range. As a consequence, estimates of juvenile production during 2014 and 2015 were among the lowest on record. These findings may be related in part to the high hatchery contribution to the population, above‐average temperatures during spawning and incubation, and superimposition rates. Management recommendations include adequate cold‐water storage in the upstream reservoir, refined spawning habitat restoration techniques in the light of superimposition rates and, on a broader system scale, actions that reduce the amount of stray hatchery Chinook salmon.     

Predation on native sculpin by exotic brown trout exceeds that by native cutthroat trout within a mountain watershed (Logan,UT, USA)

We explored potential negative effects of exotic brown trout (Salmo trutta) on native sculpin (Cottus sp.) on the Logan River, Utah, USA by (i) examining factors most strongly correlated with sculpin abundance (e.g., abiotic conditions or piscivory?), (ii) contrasting the extent of brown trout predation on sculpin with that by native cutthroat trout (Oncorhynchus clarkii utah) and (iii) estimating the number of sculpin consumed by brown trout along an elevational gradient using bioenergetics. Abundance of sculpin across reaches showed a strong (r ≥ 0.40) and significant (P < 0.05) correlation with physical variables describing width (positive) and gradient (negative), but not with abundance of piscivorous brown trout or cutthroat trout. In mainstem reaches containing sculpin, we found fish in 0% of age‐1, 10% of age‐2 and 33% of age‐3 and older brown trout diets. Approximately 81% of fish consumed by brown trout were sculpin. Despite a similar length–gape relationship for native cutthroat trout, we found only two fish (one sculpin and one unknown) in the diets of native cutthroat trout similar in size to age‐3 brown trout. Based on bioenergetics, we estimate that an average large (> 260 mm) brown trout consumes as many as 34 sculpin per year. Nevertheless, results suggest that sculpin abundance in this system is controlled by abiotic factors and not brown trout predation. Additional research is needed to better understand how piscivory influences brown trout invasion success, including in‐stream experiments exploring trophic dynamics and interactions between brown trout and native prey under different environmental conditions.