Riverine, estuarine and marine migratory behaviour and physiology of wild and hatchery-reared coho salmon Oncorhynchus kisutch (Walbaum) smolts descending the Campbell River, BC, Canada

Eighty coho salmon Oncorhynchus kisutch smolts (40 wild and 40 hatchery-reared) were surgically implanted with acoustic transmitters and released into the Quinsam River over 2 days. Differences in physiology, travel time and migratory behaviour were examined between wild and hatchery-reared fish. In addition, tagged and control fish of both wild and hatchery-reared stock were raised for 3 months following surgery to compare survival and tag retention. Detection ranges of the acoustic receivers were tested in the river, estuary and ocean in a variety of flow conditions and tide levels. Receivers were placed in the river, estuary and up to 50 km north and south from the river mouth in the marine environment. Wild smolts were significantly smaller by mass, fork length and condition factor than hatchery-reared smolts and exhibited significantly higher levels of sodium, potassium and chloride in their blood plasma than hatchery-reared smolts. The gill Na⁺K⁺-ATPase activity was also significantly higher in the wild coho smolts at the time of release. Ninety-eight per cent of wild and 80% of hatchery-reared fish survived to the estuary, 8 km downstream of the release site. No difference was found in migration speed, timing or survival between smolts released during daylight and those released after dark. Wild smolts, however, spent less time in the river and estuary, and as a result entered the ocean earlier than hatchery-reared smolts. Average marine swimming speeds for wild smolts were double those of their hatchery-reared counterparts. While hatchery smolts dispersed in both a northward and southward direction upon entering the marine environment, the majority of wild smolts travelled north from the Campbell River estuary. The wild coho salmon smolts were more physiologically fit and ready to enter sea water than the hatchery-reared smolts, and as a result had higher early survival rates and swimming speeds.     

Comparison of UV-B tolerance between wild and hatchery-reared juveniles of red sea bream (Pagrus major) and black sea bream (Acanthopagrus schlegeli)

The amount of ultraviolet (UV)-B radiation reaching the sea surface has increased due to ozone depletion. Several laboratory studies have highlighted the negative impacts of UV radiation on fish using hatchery-reared specimens. However, potential differences in UV tolerance between wild and hatchery-reared fish have been given little consideration. Wild and reared juveniles of red sea bream and black sea bream were exposed to one of four different UV-B radiation levels (1.8; 1.1; 0.4; 0?W/m²) for 4?h. Survival rate was measured every 2?h for a period of 24?h (red sea bream) or 48?h (black sea bream) following exposure. Wild and reared juvenile red sea bream were characterized by similar survival rate, with survival declining to almost 0?% 24?h after exposure at the 1.1 and 1.8?W/m² levels. In black sea bream, wild individuals showed significantly higher survival than reared fish in levels 1.1 and 1.8?W/m². Melanophore density was also measured since melanin absorbs UV radiation. Wild black sea bream showed higher melanophore density compared to reared individuals, while no such difference was observed in red sea bream. We conclude that wild black sea bream juveniles acquire higher UV tolerance partly by increasing melanophore density through exposure to UV radiation. Our results indicate that the predicted impacts of UV radiation on fish populations solely based on experimentation with hatchery-reared specimens may be overestimated for some species.     

Differences in the pattern of antipredator behaviour between hatchery‐reared and wild European sea bass juveniles

Shoals of hatchery‐reared and wild sea bass juveniles Dicentrarchus labrax were tested for differences in their antipredator responses towards a potential live predator, the eel Anguilla anguilla . Eight experimental shoals ( i.e . replicates), each composed of 15 individuals from the same stock of juveniles ( i.e . wild or hatchery), were video recorded for 5 min before and after predator exposure. A set of behavioural variables were measured during the pre‐stimulus and stimulus phases of each test and compared between the two groups of replicates. Results showed that in both hatchery‐reared and wild juveniles predator exposure elicited a significant increase in the mean level of shoal cohesiveness and mean shoal distance from the predator, and a significant decrease in the mean shoal distance from the bottom. Shoals of wild juveniles, however, aggregated more quickly and reached higher shoal cohesiveness within the first 20 s of the stimulus period than shoals of hatchery‐reared fish. During this period, the wild fish also reached the highest peak in shoal cohesiveness, which then decreased gradually towards the levels observed before predator exposure. Another component of the antipredator response, the predator inspection behaviour, was fully developed in both wild and hatchery fish. Wild fish, however, tended to inspect the predator at a closer distance than hatchery fish.     

Growth and social interactions of wild and sea-ranched brown trout and their hybrids

A laboratory study of the social behaviour and growth performance of juvenile brown trout Salmo trutta of wild and sea-ranched origin and their crosses, indicated that the social behaviour of wild and sea-ranched fish differed. Male and female parents seemed to have a different impact on the juveniles. The category having wild mother and sea-ranched father were less aggressive, less active in general, but active in feeding, and therefore had higher growth rates. This pattern arose despite that the feeding rate and the motivation to first get a food item when food was provided did not differ between the groups of fish. Wild fish tended to be most aggressive. If sea-ranched and wild fish have different intrinsic (genetically based) 'life styles', the crosses between wild and sea-ranched fish indicate that there is likelihood for an introgression of genes adapted to hatchery environment into the genetics of wild conspecifics. This is particularly serious when hatchery-reared fish invades wild populations over many years.     

Prior residence, aggression and territory acquisition in hatchery-reared and wild brown trout

In an artificial stream environment, established wild brown trout initiated 44% of the mean aggressive acts whilst hatchery-reared trout initiated 34% and introduced wild trout initiated 22%. Established wild fish maintained home stations closer to a point source of feed than did both hatchery-reared and introduced wild conspecifics. Established wild fish were the only group to show a positive mean specific growth rate during the trials. Introduced wild fish showed a slightly negative mean specific growth rate, whilst introduced hatchery-reared fish exhibited a considerable negative mean specific growth rate. These results suggest that established wild brown trout in a semi-natural stream environment display a prior-resident effect over late introductions of hatchery-reared and wild conspecifics. Introduced hatcheryreared fish were more aggressive and exhibited a lower mean specific growth rate than simultaneously stocked wild fish, suggesting that excessive expenditure of energy for unnecessary aggression may contribute to the poor survival of hatchery-reared fish after they are stocked into streams.     

Migration, growth and survival of wild and hatchery-reared anadromous Arctic charr (Salvelinus alpinus) in Finnmark, Northern Norway

Two groups of anadromous Arctic charr ( Salvelinus alpinus ) (size 200–350 mm) reared in heated water (6–12° C) under simulated natural photoperiod were individually tagged and released in spring 1988. The fish were released at two sites, in the estuary of the River Halselva and in the fjord, 2 km from the river mouth. Growth, timing of migration and survival of these hatchery-reared fish was compared to that of wild anadromous charr of the same size over a 4-year period. The hatchery-reared charr had poorer growth than the wild fish during their first year in sea water. They also resided longer in the sea and had a slightly lower survival than wild fish. During the second year, hatchery-reared charr displayed good growth, and after the third sea-season the fish were ready for slaughter at a size of approximately 800g. The results suggest that the successful development of Arctic charr ranching will be dependent upon production and release strategies that lead to improved migratory and feeding behaviour of the fish during their first season at sea.     

Trade-off between egg mass and egg number in brown trout

Individual egg mass and fecundity increased with somatic mass in first time and repeat spawning wild anadromous and freshwater resident brown trout Salmo trutta . The egg mass was larger for similar-sized trout in south (58° N) than mid Norway (63° N), whereas fecundity was higher in mid- than in south Norway, making total gonadal investment similar in the two areas. Repeat spawners had heavier eggs than similar-sized first time spawners. The egg mass of residents was c. 10% larger than that of similar-sized first time spawning anadromous trout. Common garden experiments with offspring of wild anadromous trout showed no significant correlation between egg and somatic mass in first time spawners in two of the three populations studied. In the third population, a slight positive correlation was found. Similar results were found for repeat spawners. In the three populations, fecundity increased significantly with somatic mass in both first time and repeat spawners. Wild and hatchery-reared trout showed negative correlation between egg mass and fecundity when the effect of body size was excluded, indicating a trade-off between the two parameters. In wild trout, this was caused by variation among populations, whereas in hatchery fish, within population variation was observed in egg mass over fecundity. Furthermore, the egg mass of first time and repeat spawners were positively correlated, when adjusted for fish size. Size-specific gonadal investment was significantly higher in wild anadromous than resident trout. There was no significant difference in gonadal investment between first time and repeat spawners in wild anadromous trout. However, in the hatchery-reared trout, gonadal investment was significantly higher at repeat than first time maturation. The hatchery trout did not spawn naturally, but were artificially stripped. Among populations, a part of the variation in egg mass and fecundity is phenotypically plastic, a part appears genetically determined.     

Experience and social environment influence the ability of young brown trout to forage on live novel prey

Efficient feeding is crucial for the growth, survival and reproductive success of most animals. In artificial-rearing environments, however, animals are deprived of many stimuli normally experienced in the wild, which may alter feeding behaviour, and thus influence their survival and reproductive success upon release in nature. In a laboratory experiment, we investigated the effect of hatchery rearing on the ability of brown trout, Salmo trutta, to capture and consume a novel live prey item. Hatchery-reared and wild-caught trout, originating from the same river, were fed single black crickets, either in isolation or in visual and olfactory contact with another hatchery-reared or wild-caught fish. Total consumption, time to first bite and feeding efficiency were monitored. Wild-caught trout ate more, were quicker to attack, and consumed attacked prey more efficiently than hatchery-reared fish. Food consumption and efficiency increased in both wild and hatchery-reared trout during the experiment. We propose that the differences in feeding ability between wild-caught and hatchery-reared brown trout were mainly due to differences in previous experience of feeding on live prey. Wild-caught trout tended to eat more and sooner when in visual contact with another fish than when in isolation. This trend was not seen for the hatchery-reared fish, which may be due to environmental differences between the hatchery and the natural stream. The initial inability of hatchery-reared fish to forage on live prey may reduce their success when released in the wild, especially when in competition with resident wild fish. Copyright 2001 The Association for the Study of Animal Behaviour.     

Influence of mass release of hatchery-reared Japanese flounder on the feeding and growth of wild juveniles in a nursery ground in the Japan Sea

The feeding and growth of wild juvenile Japanese flounder Paralichthys olivaceus (Temminck et Shlegel), collected after experimental mass releases of marked hatchery-reared juveniles of the same species, was examined at Wada Beach, Wakasa Bay, the Japan Sea, in 1997 and 1998. The feeding and growth of wild juveniles in the eastern area where the hatchery-reared juveniles were released and recaptured were compared with those in the western area (control area) where no hatchery-reared juveniles were recaptured. The stomach contents of both hatchery-reared and wild juveniles collected in the eastern and western areas consisted mainly of mysids in the early season (late-May to mid-June), and fish larvae in the late season (late-June to early July) in both 1997 and 1998. The stomach content index (SI) of juvenile flounder collected in the eastern and western areas showed that the feeding conditions followed similar patterns. However, the percent of empty stomachs in the wild flounder juveniles collected within 4 days after the mass release in the eastern area was as high as 10-30% in 1997 and 40% in 1998, whereas it was 0% in the western area in both years. Otolith microstructure analysis showed that the growth rate in the eastern area was not significantly different before and after release in both years. These results demonstrate that the mass release of hatchery-reared juvenile flounder temporarily affects the feeding efficiency of the cohabiting wild juveniles, but not to an extent that significantly reduces the growth rate.     

The role of experience and chemical alarm signalling in predator recognition by fathead minnows, Pimephales promelas

Young-of-the-year, predator-naive fathead minnows, Pimephales promelas , from a pikesympatric population did not respond to chemical stimuli from northern pike, Esox Indus , while wild-caught fish of the same age and size did. These results suggest that chemical predator recognition is a result of previous experience and not genetic factors, Wild young-of-the-year minnows responded to pike odour with a response intensity that was similar to that of older fish, demonstrating that the ability to recognize predators is learned within the first year. The intensity of response of wild minnows which had been maintained in a predator free environment for 1 year was similar to that of recently caught minnows of the same age, suggesting that reinforcement was not required for predator recognition to be retained. Naive minnows that were exposed simultaneously to chemical stimuli from pike (a neutral stimulus) and minnow alarm substance exhibited a fright response upon subsequent exposure to the pike stimulus alone. Predator-naive minnows exposed simultaneously to chemical stimuli from pike and glass-distilled water did not exhibit a fright response to the pike stimulus alone. These results demonstrate that fathead minnows can acquire predator recognition through releaserinduced recognition learning, thus confirming a known mechanism through which alarm substance may benefit the receivers of an alarm signal.     

Hatchery-reared fish have less consistent behavioral pattern compared to wild individuals,exemplified by red tilefish studied using video observation and acoustic telemetry tracking

The behavior of wild and hatchery-reared red tilefish Branchiostegus japonicus was analyzed using two different methods: video observation and acoustic telemetry tracking. In the laboratory, digging and swimming activity of seven wild and five hatchery-reared fish were recorded for 2–4 days in an experimental aquarium and related to changes in light intensity. The activity of wild individuals increased with light intensity, while hatchery-reared individuals were active during both day and night. In the field, 18 wild and 9 hatchery-reared fish were released and tracked using an acoustic telemetry system during the winter and the summer in Maizuru Bay, Kyoto (Japan). Seven wild and three hatchery-reared fish settled within 2 km of the release point for 21–200 days whilst 11 wild and 6 hatchery-reared fish were detected in the central part of the bay within 13 days. The settled fish demonstrated a diel pattern of behavior; most of them probably stayed outside their burrows in the daytime but inside their burrows during the nighttime. During the winter wild fish tended to stay near the release site for longer periods than the hatchery-reared fish. However, in summer the hatchery-reared fish tended to remain close to the release site for longer than the wild fish. Based on these two experiments, we suggest that the activity of wild red tilefish respond to changes in low levels of light intensity (0–0.1 μmol m⁻² s⁻¹) at dawn and display a diel behavior. Hatchery-reared fish may differ from wild fish with regard to their behavior, response to water temperature and light conditions. Electronic supplementary material Supplementary material is available for this article at and accessible for authorized users     

Differences in Lateral Line Morphology between Hatchery- and Wild-Origin Steelhead

Despite identification of multiple factors mediating salmon survival, significant disparities in survival-to-adulthood among hatchery- versus wild-origin juveniles persist. In the present report, we explore the hypothesis that hatchery-reared juveniles might exhibit morphological defects in vulnerable mechanosensory systems prior to release from the hatchery, potentiating reduced survival after release. Juvenile steelhead (Oncorhynchus mykiss) from two different hatcheries were compared to wild-origin juveniles on several morphological traits including lateral line structure, otolith composition (a proxy for auditory function), and brain weight. Wild juveniles were found to possess significantly more superficial lateral line neuromasts than hatchery-reared juveniles, although the number of hair cells within individual neuromasts was not significantly different across groups. Wild juveniles were also found to possess primarily normal, aragonite-containing otoliths, while hatchery-reared juveniles possessed a high proportion of crystallized (vaterite) otoliths. Finally, wild juveniles were found to have significantly larger brains than hatchery-reared juveniles. These differences together predict reduced sensitivity to biologically important hydrodynamic and acoustic signals from natural biotic (predator, prey, conspecific) and abiotic (turbulent flow, current) sources among hatchery-reared steelhead, in turn predicting reduced survival fitness after release. Physiological and behavioral studies are required to establish the functional significance of these morphological differences.     

Predator avoidance behaviour in wild and hatchery‐reared brown trout: the role of experience and domestication

Juvenile brown trout Salmo trutta from natural populations reacted to the presence of piscivorous brown trout by increasing the use of refuges. In contrast, second‐generation hatchery fish and the offspring of wild fish raised under hatchery conditions were insensitive to predation risk. The diel pattern of activity also differed between wild and hatchery brown trout. Second‐generation hatchery fish were predominantly active during daytime regardless of risk levels. Wild fish, however, showed a shift towards nocturnal activity in the presence of predators. These findings emphasize the potential role of domestication in weakening behavioural defences. They support the idea that the behavioural divergence between wild and domesticated individuals can arise from a process of direct or indirect selection on reduced responsiveness to predation risk, or as a lack of previous experience with predators.     

Morphometric comparison and differentiation between artificially-released and wild red sea bream (Pagrus major)

The objective of this study is to develop a method of differentiation of hatchery-reared and artificially-released red sea bream (Pagrus major) from wild fish, based upon their morphometric differences. Morphometric measurements were done on fork length and 14 other characters. Among these charac)ters, significant differences between hatchery-reared and wild red sea bream were observed in body height, height at eye, eye diameter and upper jaw length. Discriminant functions were effective in differentiating artificial fish from wild fish.     

Brown trout Salmo trutta express different morphometrics due to divergence in the rearing environment

Variation in the body morphology of juvenile brown trout Salmo trutta was studied for both wild and hatchery-reared individuals from the same gene pool. The thin-plate spline (TPS) method by pointing landmarks was used to characterize juveniles from their natal environment, the River Kuusinkijoki in eastern Finland, and from wild parents raised in a hatchery environment. Differences were found in the morphometrics of juvenile S. trutta from the two different environments. Wild S. trutta are characterized by a longer head and shorter anterior part of the trunk compared to hatchery S. trutta. Stocked to their natal river with wild S. trutta, the hatchery fish became characterized with more similar morphometrics to their wild counterparts. The characteristics of the body form are explained by the differing environmental conditions in the wild and hatchery. It is concluded that a simple hatchery environment may delay the development of morphological characteristics important in a natural river environment.     

Adaptive winter survival strategies: defended energy levels in juvenile Atlantic salmon along a latitudinal gradient

Current knowledge suggests that patterns of energy storage and depletion in animals are governed by behavioural trade-offs between risks associated with feeding and future energy demands. However, the length of adverse periods varies over geographical or climatic gradients. To explore the potential for genotypic sources of variation in behavioural trade-offs, we compared the winter energy-depletion patterns among 13 wild populations of juvenile Atlantic salmon (Salmo salar L.) along a latitudinal gradient (58–70°N) and performed common-environment experiments of energy-state-dependent feeding. In the wild, winter lipid-depletion rates were lower for northern than for southern populations. The variation in spring lipid levels among the population was lower than autumn variation, with storage lipid levels clustered close to critical limits for survival. In semi-natural stream channels with natural food supply, hatchery-reared fish originating from northern populations showed a positive scaling of feeding activity with decreasing energy levels, whereas southern populations did not. In conclusion, juvenile Atlantic salmon from northern populations defend their energy levels more strongly than fish from southern populations. Adaptive variation in feeding activity appears important for this difference. Thus, the present study shows a link between geographical patterns in storage energy trajectories and adaptive differences in state-dependent feeding motivation.     

Behavioural Stress Responses Predict Environmental Perception in European Sea Bass (Dicentrarchus labrax)

Individual variation in the response to environmental challenges depends partly on innate reaction norms, partly on experience-based cognitive/emotional evaluations that individuals make of the situation. The goal of this study was to investigate whether pre-existing differences in behaviour predict the outcome of such assessment of environmental cues, using a conditioned place preference/avoidance (CPP/CPA) paradigm. A comparative vertebrate model (European sea bass, Dicentrarchus labrax) was used, and ninety juvenile individuals were initially screened for behavioural reactivity using a net restraining test. Thereafter each individual was tested in a choice tank using net chasing as aversive stimulus or exposure to familiar conspecifics as appetitive stimulus in the preferred or non preferred side respectively (called hereafter stimulation side). Locomotor behaviour (i.e. time spent, distance travelled and swimming speed in each tank side) of each individual was recorded and analysed with video software. The results showed that fish which were previously exposed to appetitive stimulus increased significantly the time spent on the stimulation side, while aversive stimulus led to a strong decrease in time spent on the stimulation side. Moreover, this study showed clearly that proactive fish were characterised by a stronger preference for the social stimulus and when placed in a putative aversive environment showed a lower physiological stress responses than reactive fish. In conclusion, this study showed for the first time in sea bass, that the CPP/CPA paradigm can be used to assess the valence (positive vs. negative) that fish attribute to different stimuli and that individual behavioural traits is predictive of how stimuli are perceived and thus of the magnitude of preference or avoidance behaviour.     

Do hatchery-reared sea urchins pose a threat to genetic diversity in wild populations?

In salmonids, the release of hatchery-reared fish has been shown to cause irreversible genetic impacts on wild populations. However, although responsible practices for producing and releasing genetically diverse, hatchery-reared juveniles have been published widely, they are rarely implemented. Here, we investigated genetic differences between wild and early-generation hatchery-reared populations of the purple sea urchin Paracentrotus lividus (a commercially important species in Europe) to assess whether hatcheries were able to maintain natural levels of genetic diversity. To test the hypothesis that hatchery rearing would cause bottleneck effects (that is, a substantial reduction in genetic diversity and differentiation from wild populations), we compared the levels and patterns of genetic variation between two hatcheries and four nearby wild populations, using samples from both Spain and Ireland. We found that hatchery-reared populations were less diverse and had diverged significantly from the wild populations, with a very small effective population size and a high degree of relatedness between individuals. These results raise a number of concerns about the genetic impacts of their release into wild populations, particularly when such a degree of differentiation can occur in a single generation of hatchery rearing. Consequently, we suggest that caution should be taken when using hatchery-reared individuals to augment fisheries, even for marine species with high dispersal capacity, and we provide some recommendations to improve hatchery rearing and release practices. Our results further highlight the need to consider the genetic risks of releasing hatchery-reared juveniles into the wild during the establishment of restocking, stock enhancement and sea ranching programs.     

Effects of Emergence Time and Early Social Rearing Environment on Behaviour of Atlantic Salmon: Consequences for Juvenile Fitness and Smolt Migration

Consistent individual differences in behaviour have been well documented in a variety of animal taxa, but surprisingly little is known about the fitness and life-history consequences of such individual variation. In wild salmonids, the timing of fry emergence from gravel spawning nests has been suggested to be coupled with individual behavioural traits. Here, we further investigate the link between timing of spawning nest emergence and behaviour of Atlantic salmon (Salmo salar), test effects of social rearing environment on behavioural traits in fish with different emergence times, and assess whether behavioural traits measured in the laboratory predict growth, survival, and migration status in the wild. Atlantic salmon fry were sorted with respect to emergence time from artificial spawning nest into three groups: early, intermediate, and late. These emergence groups were hatchery-reared separately or in co-culture for four months to test effects of social rearing environment on behavioural traits. Twenty fish from each of the six treatment groups were then subjected to three individual-based behavioural tests: basal locomotor activity, boldness, and escape response. Following behavioural characterization, the fish were released into a near-natural experimental stream. Results showed differences in escape behaviour between emergence groups in a net restraining test, but the social rearing environment did not affect individual behavioural expression. Emergence time and social environment had no significant effects on survival, growth, and migration status in the stream, although migration propensity was 1.4 to 1.9 times higher for early emerging individuals that were reared separately. In addition, despite individuals showing considerable variation in behaviour across treatment groups, this was not translated into differences in growth, survival, and migration status. Hence, our study adds to the view that fitness (i.e., growth and survival) and life-history predictions from laboratory measures of behaviour should be made with caution and ideally tested in nature.     

Adaptive trade-offs in juvenile salmonid metabolism associated with habitat partitioning between coho salmon and steelhead trout in coastal streams

1. Adaptive trade-offs are fundamental to the evolution of diversity and the coexistence of similar taxa and occur when complimentary combinations of traits maximize efficiency of resource exploitation or survival at different points on environmental gradients. 2. Standard metabolic rate (SMR) is a key physiological trait that reflects adaptations to baseline metabolic performance, whereas active metabolism reflects adaptations to variable metabolic output associated with performance related to foraging, predator avoidance, aggressive interactions or migratory movements. Benefits of high SMR and active metabolism may change along a resource (productivity) gradient, indicating that a trade-off exists among active metabolism, resting metabolism and energy intake. 3. We measured and compared SMR, maximal metabolic rate (MMR), aerobic scope (AS), swim performance (UCrit) and growth of juvenile hatchery and wild steelhead and coho salmon held on high- and low-food rations in order to better understand the potential significance of variation in SMR to growth, differentiation between species, and patterns of habitat use along a productivity gradient. 4. We found that differences in SMR, MMR, AS, swim performance and growth rate between steelhead trout and coho salmon were reduced in hatchery-reared fish compared with wild fish. Wild steelhead had a higher MMR, AS, swim performance and growth rate than wild coho, but adaptations between species do not appear to involve differences in SMR or to trade-off increased growth rate against lower swim performance, as commonly observed for high-growth strains. Instead, we hypothesize that wild steelhead may be trading off higher growth rate for lower food consumption efficiency, similar to strategies adopted by anadromous vs. resident brook trout and Atlantic salmon vs. brook trout. This highlights potential differences in food consumption and digestion strategies as cryptic adaptations ecologically differentiating salmonid species. 5. We hypothesize that divergent digestive strategies, which are common and well documented among terrestrial vertebrates, may be an important but overlooked aspect of adaptive strategies of juvenile salmonids, and fish in general.