The effects of prior residence on behavior and growth rates in juvenile Atlantic salmon (Salmo salar)

It is well documented that prior residence confers advantagesin territorial disputes, but its impact on other aspects ofbehavior and fitness is less understood. We tested how priorresidence influences the subsequent feeding behavior and growthperformance of dispersing Atlantic salmon fry (Salmo salar)using experimental manipulations of residence in a seminaturalstream tank. In replicated trials, groups of seven "primary"fish were released into the stream tank 3 days ahead of seven"secondary" fish. Standardized behavioral observations were madeon each fish over the following 14 days, after which all fishwere removed and measured. Primaries and secondaries were initiallythe same size and body condition and exhibited the same degreeof site fidelity. However, primaries darted higher into thewater column to intercept prey items, fed at a higher rate,and subsequently grew faster. Larger fish (in terms of body length)tended to be more dominant, and dominants grew faster than subordinates.However, there was no difference in dominance between primaries andsecondaries. These results suggest that the well-documentedadvantage of early-emerging salmon fry over late-emerging frycannot be completely attributed to intrinsic differences andthat the advantage is partly mediated via a prior residenceeffect. Furthermore, prior residents gain foraging advantageswithout necessarily becoming more dominant.     

Early predictors of life-history events: the link between first feeding date, dominance and seaward migration in Atlantic salmon, Salmo salar L.

Previous studies have suggested that the earliest fry to emerge from a salmonid redd may have an advantage in the subsequent competition for feeding sites, partly through a 'prior residence' effect. Here we examined whether there was any relationship between the relative date of first feeding and subsequent dominance status and growth in a sibling group of Atlantic salmon, Salmo salar L., fry. Earlier-feeding fry were dominant over their later-feeding siblings (controlling for prior residence), despite not being any larger. However, these early fish soon established and then maintained a size advantage. This led to an increased probability of early-feeding fish migrating to sea at age 1 year (rather than 2 or more). Thus a difference of less than 1 week in the relative timing of first feeding can translate into a year's difference in the timing of migration.     

Effects of species, culture history, size and residency on relative competitive ability of salmonids

The relative competitive ability of juvenile farm and wild salmonids was investigated to provide insight into the potential effects of introduction of cultured salmon on wild Pacific salmonid ( Oncorhynchus ) species. Aquarium experiments involving equal contests ( i.e. size matched, simultaneously introduced individuals) indicated that two wild coho salmon Oncorhynchus kisutch populations were competitively equal to a farm coho salmon population. In equal contests between farm Atlantic salmon Salmo salar (Mowi strain) and these wild coho salmon populations or coastal cutthroat trout Oncorhynchus clarki clarki , Atlantic salmon were subordinate in all cases. When Atlantic salmon were given a residence advantage, however, they were competitively equal to both wild coho salmon populations, but remained subordinate to coastal cutthroat trout. When Atlantic salmon were given a 10–30% length advantage, they were competitively equal to one wild coho salmon population but remained subordinate to the other. In equal contests in semi-natural stream channels, both wild coho and farm Atlantic salmon grew significantly more in the presence of the other species than when alone. It appears that coho salmon obtain additional food ration by out competing Atlantic salmon, whereas Atlantic salmon were stimulated to feed more in the presence of coho salmon competitors. These results suggest that wild coho salmon and cutthroat trout should out compete farm Atlantic salmon of a similar size in nature. As the relative competitive ability of Atlantic salmon improves when they have a size and residence advantage, should feral populations become established, they may exist on a more equal competitive footing owing to the long freshwater residence of Atlantic salmon.     

Fish may fight rather than feed in a novel environment: metabolic rate and feeding motivation in juvenile Atlantic salmon

This study tested the hypothesis that juvenile Atlantic salmon Salmo salar with a high resting metabolic rate and probability of dominance will also have a correspondingly higher feeding motivation in a novel environment to offset their greater costs of maintenance. The opposite was found to be the case: Atlantic salmon with a high standard metabolic rate had a slightly but significantly lower feeding motivation. It is hypothesized that Atlantic salmon with higher maintenance costs opt to be more aggressive at the expense of the costly activities associated with feeding, since elevated aggression in a new habitat is a more successful strategy for acquiring a feeding territory (and hence sustaining food intake in the long-term) than a high feeding motivation alone.     

The performance advantage of a high resting metabolic rate in juvenile salmon is habitat dependent

1.?Basal levels of metabolism vary significantly among individuals in many taxa, but the effects of this on fitness are generally unknown. Resting metabolic rate (RMR) in juvenile salmon and trout is positively related to dominance status and ability to obtain a feeding territory, but it is not clear how this translates into performance in natural conditions. 2.?The relationships between RMR, dominance, territoriality and growth rates of yearling Atlantic salmon Salmo salar were examined in relation to predictability in food supply and habitat complexity, using replicate sections of a large-scale controlled semi-natural stream. 3.?Estimated RMR was a strong predictor of dominance, and under conditions of a predictable food supply in a structurally simple habitat, high estimated RMR fish obtained the best feeding territories and grew faster. 4.?When the spatial distribution of food was made less predictable, dominant (high estimated RMR) fish were still able to occupy the most profitable feeding locations by periodically moving location to track the changes in food availability, but RMR was no longer a predictor of growth rate. Moreover, when a less predictable food supply was combined with a visually more complex (and realistic) habitat, fish were unable to track changes in food availability, grew more slowly and exhibited greater site fidelity, and there were no relationships between estimated RMR and quality of occupied territory or growth rate. 5.?The relative benefit of RMR is thus context dependent, depending on both habitat complexity and the predictability of the food supply. Higher habitat complexity and lower food predictability decrease the performance advantages associated with a high RMR.     

Competition between brown trout and Atlantic salmon parr over pool refuges during rapid dewatering

Variations in distributions and behaviours of Atlantic salmon Salmo salar in allopatry (homogeneous) and in sympatry with brown trout Salmo trutta (mixed) were observed before, during and after 2 day periods of dewatering in a large glass-sided indoor stream at densities typical of Scottish upland streams. Brown trout utilized pools more than Atlantic salmon at normal flows and in both species the majority of fishes moved into pools during dewatering. There was no significant effect of brown trout, which was the more dominant species, on the overall ability of Atlantic salmon to use pool habitat as a refuge during dewatering. Within mixed and homogeneous groups, average feeding levels decreased during dewatering. The highest ranking fish, which was always a brown trout in mixed groups, predominantly monopolized the pool and other individuals in pools adopted a more cryptic, stationary behaviour. Dewatering effectively increased local population density with the result that dominance status became much more important in maintaining food intake, and polarization between the top ranking fish and others increased. During the first day of dewatering, there was extreme behavioural polarization such that the dominant fish exhibited most aggression and least feeding within the group. Among dominant fish on the second day of dewatering, aggression had largely abated and feeding had returned to pretreatment levels despite the reduced average feeding within the group. The main difference between mixed and homogeneous groups was in the behaviour of the most dominant Atlantic salmon, which was near-despotic in allopatry and subordinate to brown trout in sympatry.     

Surface feeding and aggressive behaviour of diploid and triploid brown trout Salmo trutta during allopatric pair‐wise matchings

Diploid and triploid brown trout Salmo trutta were acclimated for 6 weeks on two feeding regimes (floating and sinking). Thereafter, aggression and surface feeding response were compared between pairs of all diploid, all triploid and diploid and triploid S. trutta in an experimental stream. In each pair‐wise matching, fish of similar size were placed in allopatry and rank was determined by the total number of aggressive interactions recorded. Dominant individuals initiated more aggression than subordinates, spent more time defending a territory and positioned themselves closer to the surface food source (Gammarus pulex), whereas subordinates occupied the peripheries. In cross ploidy trials, diploid S. trutta were more aggressive than triploid, and dominated their sibling when placed in pair‐wise matchings. Surface feeding, however, did not differ statistically between ploidy irrespective of feeding regime. Triploids adopted a sneak feeding strategy while diploids expended more time defending a territory. In addition, we also tested whether triploids exhibit a similar social dominance to diploids when placed in allopatry. Although aggression was lower in triploid pairs than in the diploid and triploid pairs, a dominance hierarchy was also observed between individuals of the same ploidy. Dominant triploid fish were more aggressive and consumed more feed items than subordinate individuals. Subordinate fish displayed a darker colour index than dominant fish suggesting increased stress levels. Dominant triploid fish, however, appeared to be more tolerant of subordinate individuals and did not display the same degree of invasive aggression as seen in the diploid and diploid or diploid and triploid matchings. These novel findings suggest that sterile triploid S. trutta feed similarly but are less aggressive than diploid trout. Future studies should determine the habitat choice of triploid S. trutta after release and the interaction between wild fish and triploids during the breeding season prior to utilization of triploids as an alternative management strategy within freshwater fisheries.     

Stressed mothers - troubled offspring: a study of behavioural maternal effects in farmed Salmo salar

Mature female Atlantic salmon Salmo salar were given intraperitoneal cortisol implants 1 week prior to stripping to examine the influence of simulated maternal stress on offspring boldness and social dominance. Behavioural tests originally designed to investigate stress responsiveness and coping styles in salmonids (i.e. feeding in isolation, dominance tests and acute confinement) were carried out on the offspring 1·5 years after hatching. In the feeding test, there were no differences between the two treatment groups in total feeding score or number of pellets eaten, but offspring from the cortisol-implanted females made more unsuccessful feeding attempts than offspring from control females. In dominance tests, there was no difference between controls and cortisol-treated fish regarding propensity to become socially dominant. A higher proportion of individuals with bite marks, however, was observed in the cortisol group when compared to controls. Cortisol-treated offspring that gained dominant rank in the dominance tests performed more aggressive acts after stable dominance-subordinate relationships were established compared to control winners. During acute confinement stress, offspring from cortisol-implanted females showed a reduction in the proportion of time they were moving compared to the controls. These results indicate that the maternal endocrine state at spawning affects several aspects of progeny behaviour potentially related to subsequent success and survival in farmed S. salar.     

Interspecific habitat associations of juvenile salmonids in Lake Ontario tributaries: implications for Atlantic salmon restoration

Diel variation in habitat use of subyearling Chinook salmon (Oncorhynchus tshawytscha), subyearling coho salmon (O. kisutch), yearling steelhead (O. mykiss), and yearling Atlantic salmon (Salmo salar) was examined during the spring in two tributaries of Lake Ontario. A total of 1318 habitat observations were made on juvenile salmonids including 367 on steelhead, 351 on Chinook salmon, 333 on Atlantic salmon, and 261 on coho salmon. Steelhead exhibited the most diel variation in habitat use and Chinook the least. Juvenile salmonids were generally associated with more cover and larger substrate during the day in both streams. Interspecific differences in habitat use in both streams occurred with Atlantic salmon (fast velocities) and coho salmon (pools) using the least similar habitat. Chinook salmon and Atlantic salmon used similar habitat in both streams. These findings should help guide future management actions specific to habitat protection and restoration of Atlantic salmon in Lake Ontario tributaries.     

Visual isolation furthers access to drift‐feeding positions for subordinate juvenile brown trout (Salmo trutta) in dominance hierarchies

Juvenile salmonids are visual predators that primarily feed on drifting invertebrates and compete for suitable feeding positions in swift water. We used an outdoor experimental stream to test the effect of visual isolation on agonistic interactions and habitat use by age‐1 brown trout ( Salmo trutta ) in riffle‐pool sections. We hypothesized that dominant fish defend suitable feeding positions in riffle and that visual obstruction between individuals enhance access to riffle for subordinates. Groups of juveniles, caught in the wild, were stocked in high and low visibility treatments. Visual isolation was manipulated by placing dark plastic ribbons or opaque Plexiglas boards onto the substrate of riffles. As expected, dominant fish held profitable positions in riffle and the proportion of fish in riffle significantly increased in presence of artificial structures. In high visibility treatment, the dominant fish despotically excluded subordinates from the riffle. In low visibility treatment, the proportion of fish that foraged on benthic prey in the pool and the number of major aggressive acts (chase, nip) decreased. Our results support the hypothesis that screening effect of physical structures in the water column loosen resource monopolization in dominance hierarchies of juvenile salmonids.     

Juvenile salmon in estuaries: comparisons between North American Atlantic and Pacific salmon populations

All anadromous fishes, including juvenile salmon, encounter estuarine habitats as they transition from riverine to marine environments. We compare the estuarine use between juvenile Atlantic salmon (Salmo salar) in the Penobscot River estuary and Pacific salmon (Oncorhynchus spp.) in the Columbia River estuary. Both estuaries have been degraded by anthropogenic activities. Atlantic and Pacific salmon populations in both basins rely heavily on hatchery inputs for persistence. Pacific salmon, as a group, represent a continuum of estuarine use, from species that move through rapidly to those that make extensive use of estuarine habitats. While Atlantic salmon estuarine use is predominantly similar to rapidly moving Pacific salmon, they can exhibit nearly the entire range of Pacific salmon estuarine use. Both slow and rapidly migrating Atlantic and Pacific salmon actively feed in estuarine environments, consuming insect and invertebrate prey. Interactions between juvenile salmon and estuarine fish communities are poorly understood in both estuaries, although they experience similar avian and marine mammal predators. Estuaries are clearly important for Atlantic and Pacific salmon, yet our understanding of this use is currently insufficient to make informed judgments about habitat quality or overall estuary health. This review of salmonid migration through and residency within estuaries identifies actions that could hasten restoration of both Atlantic and Pacific salmon populations.     

Dominance hierarchies among communally held juvenile lobsters,Homarus americanus

An investigation was conducted to determine the time for establishment of dominance hierarchies, their stability, and their relationship to molting and death of cultured juvenile lobsters, Homarus americanus. Agonistic displays between communally held animals were monitored daily. Interactions between all possible pairs of animals within each group were used to determine formation of a dominance hierarchy. A decrease in the frequency of agonistic interactions was recorded after the first day of the experiment. A stable four‐rank hierarchy was established within 2–3 days. When the dominant animal was molting, the subordinates displayed aggressively toward all members of the group but remained submissive to the dominant. When the dominant animal died, stability was lost and aggressive interactions among subordinates established a new ranking. Molting subordinates were highly susceptible to injury and death. Loss of subordinates did not affect the ranking of individuals.     

Habitat use and fish activity of landlocked Atlantic salmon and brook charr in a newly developed habitat compensation facility

Degradation and destruction of valuable spawning and rearing habitat due to anthropogenic changes (e.g., flow modification and channelisation) is known to have dramatic impacts on fish populations. To compensate for habitat losses due to hydropower development, an artificial fluvial habitat channel (‘Compensation Creek’) was constructed in south-central Newfoundland, Canada. The creek was designed to include appropriate habitat features for the two dominant salmonid fish species, landlocked Atlantic salmon (Salmo salar L.) and brook charr (Salvenius fontinalis Mitchell). The study examines the habitat use of landlocked Atlantic salmon and brook charr in the Compensation Creek using electromyogram (EMG) radio telemetry. Ten landlocked Atlantic salmon and eight brook charr were captured and tagged with EMG transmitters. In laboratory swimming experiments, the EMG values were calibrated against swimming speed. Fish were then released in the Compensation Creek and tracked on a daily basis. The results show that (1) during residence in the creek, both species used preferentially the habitat features designed to match their rearing habitat preferences, and (2) swimming speed did not vary among habitat types for either species.     

Establishment of a free-ranging colony of stumptail macaques (Macaca arctoides): Social relations I

Data on the social behaviour of a free-ranging troop of Stumptail macaques are reported for a period starting of August 15th to November 7th of 1974. Positive interactions take a greater part in the daily life of the troop than negative interactions. Individuals order themselves in an essentially linear dominance hierarchy and offspring rank immediately below their mother. Agonism and displacement are good predictors of dominance rank. Dominant individuals are a strong source of attraction for subordinates and the alpha male is a powerful social magnet for immatures. Grooming was found to be correlated with dominance rank and individuals holding adjacent ranks tend to interact positively with one another. To gain access to dominant females, subordinates direct a great deal of positive behaviour to the female’s youngest offspring. As social distance decreases the selection of a social partner becomes more strict and tends toward adjacency. However, factors such as motherhood, genealogical ties, and special relationships with non adjacent individuals, can override the tendency. The direction and quantity of male-care displayed by the adult male and by juvenile males toward younger individuals was found to be influenced by the troop’s dominance patterns. Sex differences were found to exist in male-care received and a negative association was found with the age of the receiver. Adult females are the main regulators of social relations and their rank and friendships provide their offspring with the social format, before puberty, in which they develop socially. The alpha male is the group protector, the performer of policing, an important element in the immatures’ social development, and a powerful source of attraction for all troop members.     

Small Subordinate Male Advantage in the Zebrafish

Dominance is an important determinant of reproductive success in many species, and size is usually an indicator of dominance status, with larger, dominant individuals physically and physiologically preventing smaller subordinates from mating. However, small size may be advantageous in some mating contexts because enhanced manoeuvrability enables males to get closer to females during mating. Here, we determined the paternity success and testes size of dominant and subordinate male zebrafish (Danio rerio), in pairs that controlled for social status. There was no statistical difference in both body size and testes size between dominant and subordinate males. Dominant males sired significantly more offspring than subordinates, but when subordinates were small, they had a greater share of the paternity than larger subordinates. Small male advantage may be one mechanism by which variation in body size is maintained in this species.     

Effects of location of food delivery and social status on foraging-site selection by juvenile Atlantic salmon

Synopsis Experiments were conducted to investigate the response of juvenile Atlantic salmon,Salmo salar, to changes in the location at which food entered a stream tank. Experience with the location of food input into the system significantly influenced foraging-site selection by the dominant fish. Dominant fish changed location of their foraging site in response to a change in the location of food input, and occupied, aggressively defending, sites just downstream of the location of food introduction. In contrast, subordinate fish occupied foraging sites at the downstream end of the stream tank, regardless of location of food input. As a result of both site selection and social status, dominant fish captured significantly more prey than subordinates. Our results support the contention that salmonids choose foraging sites to maximize foraging opportunities. Our results also provide a possible explanation for the use of atypical foraging sites by individual fish within their home range over the course of a single day, as observed in a number of salmonid species in the field.     

Endocrine mediated phenotypic plasticity: Condition-dependent effects of juvenile hormone on dominance and fertility of wasp queens

There has been increasing interest in the mechanisms that mediate behavioral and physiological plasticity across individuals with similar genotypes. Some of the most dramatic plasticity is found within and between social insect castes. For example, Polistes wasp queens can nest alone, dominate a group of cooperative queens, or act as worker-like subordinates who rarely reproduce. Previous work suggests that condition-dependent endocrine responses may play a role in plasticity between castes in the hymenoptera. Here, we test whether condition-dependent endocrine responses influence plasticity within castes in the wasp Polistes dominulus. We experimentally manipulate juvenile hormone (JH) titers in nest-founding queens and assess whether JH mediates variation in behavior and physiology. JH generally increased dominance and fertility of queens, but JH's effects were not uniform across individuals. JH had a stronger effect on the dominance and fertility of large individuals and individuals with facial patterns advertising high quality than on the dominance and fertility of small individuals and those advertising low quality. These results demonstrate that JH has condition-dependent effects. As such, they clarify how JH can mediate different behaviors in well nourished queens and poorly nourished workers. Many Polistes queens nest cooperatively with other queens, so condition-dependent hormonal responses provide a mechanism for queens to adaptively allocate energy based on their probability of successfully becoming the dominant queen. Research on the endocrine basis of plasticity often focuses on variation in endocrine titers alone. However, differential endocrine responses are likely to be a widespread mechanism mediating behavioral and physiological plasticity.     

An evaluation of the quality of fall-fingerling Atlantic salmon (Salmo salar L.) released to natural lacustrine nursery areas in Newfoundland,Canada

Results of experiments on controlled releases of fall-fingerling Atlantic salmon to lacustrine habitats in Newfoundland are reviewed in terms of survival during freshwater residence and smolt yield per unit of standing water habitat. Average survival from parr to smolt in three consecutive annual releases was 12.9% and we conclude that the quality of fall-fingerling salmon, produced from our semi-natural rearing techniques, was satisfactory. Year-class survival decreased progressively with consecutive releases. Rate of increase in juvenile salmon biomass remained stable over the duration of the study, suggesting that the average annual biomass of 1.6 kg ha^–1 of juvenile salmon released did not exceed habitat carrying capacity. Precocity did not appear to represent a significant deterrent to fall-fingerling survival. Annual smolt yield (by year-class) from stocking ranged from 53 to 130 ha^–1 (mean = 77 smolts ha^–1).     

Hatching Time and Alevin Growth Prior to the Onset of Exogenous Feeding in Farmed,Wild and Hybrid Norwegian Atlantic Salmon

The onset of exogenous feeding, when juveniles emerge from the gravel, is a critical event for salmonids where early emergence and large size provide a competitive advantage in the wild. Studying 131 farmed, hybrid and wild Norwegian Atlantic salmon families, originating from four wild populations and two commercial strains, we investigated whether approximately 10 generations of selection for faster growth has also resulted in increased somatic growth prior to the onset of exogenous feeding. In addition, we tested whether relaxed selection in farms has allowed for alterations in hatching time between farmed and wild salmon. Across three cohorts, wild salmon families hatched earlier than farmed salmon families, while hybrid families displayed intermediate hatching times. While the observed differences were small, i.e., 1–15 degree-days (0–3 days, as water temperatures were c. 5–6°C), these data suggest additive genetic variation for hatching time. Alevin length prior to exogenous feeding was positively related to egg size. After removal of egg size effects, no systematic differences in alevin length were observed between the wild and farmed salmon families. While these results indicate additive genetic variation for egg development timing, and wild salmon families consistently hatched earlier than farmed salmon families, these differences were so small they are unlikely to significantly influence early life history competition of farmed and wild salmon in the natural environment. This is especially the case given that the timing of spawning among females can vary by several weeks in some rivers. The general lack of difference in size between farmed and wild alevins, strongly suggest that the documented differences in somatic growth rate between wild and farmed Norwegian Atlantic salmon under hatchery conditions are first detectable after the onset of exogenous feeding.     

Diet,feeding patterns,and prey selection of subyearling Atlantic salmon (Salmo salar) and subyearling chinook salmon (Oncorhynchus tshawytscha) in a tributary of Lake Ontario

Since juvenile Atlantic salmon (Salmo salar) and Chinook salmon (Oncorhynchus tshawytscha) occupy a similar habitat in Lake Ontario tributaries, we sought to determine the degree of diet similarity between these species in order to assess the potential for interspecific competition. Atlantic salmon, an historically important but currently extirpated component of the Lake Ontario fish community, are the focus of a bi‐national restoration effort. Presently this effort includes the release of hatchery produced juvenile Atlantic salmon in Lake Ontario tributaries. These same tributaries support substantial numbers of naturally reproduced juvenile Pacific salmonids including Chinook salmon. Subyearling Atlantic salmon and subyearling Chinook salmon had significantly different diets during each of the three time periods examined. Atlantic salmon fed slightly more from the benthos than from the drift and consumed mainly chirononmids (47.0%) and ephemeropterans (21.1%). The diet of subyearling Chinook salmon was more closely associated with the drift and consisted mainly of chironomids (60.2%) and terrestrial invertebrates (16.0%). Low diet similarity between subyearling Atlantic salmon and subyearling Chinook salmon likely minimizes competitive interactions for food between these species in Lake Ontario tributaries. However, the availability of small prey such as chironomids which comprise over 50% of the diet of each species, soon after emergence, could constitute a short term resource limitation. To our knowledge this is the first study of interspecific diet associations between these two important salmonid species.