The disruption of dominance hierarchies by a non-native species: an individual-based analysis

We studied the effects of the exotic rainbow trout (Oncorhynchus mykiss) on the performance and the dominance hierarchy of native Atlantic salmon (Salmo salar) at the group and individual level using laboratory and semi-natural experiments. At the group level, we compared the effects of interspecific and intraspecific competition (substitutive and additive design) on behavioural responses and growth of young-of-the-year Atlantic salmon. At the individual level, the same design was used to evaluate: (1) the temporal consistency of behavioural responses, dominance hierarchy and growth rate of Atlantic salmon; (2) the pattern of correlations between behaviours; and (3) the relationship between individual growth rate and behaviour. In the laboratory, group-level analyses revealed a weak but similar effect of rainbow trout and intraspecific competition on the behaviour and growth of Atlantic salmon. In contrast, individual-based analyses demonstrated that rainbow trout (but not intraspecific competition) strongly affected behavioural strategy, dominance hierarchy and growth trajectory of individual Atlantic salmon. Specifically, behaviours, dominance status and growth rate of salmon were temporally consistent in the intraspecific environment, while these patterns were disrupted when rainbow trout were present. Similarly, we found that rainbow trout strongly affected behavioural correlations and the relationships between individual growth rate and behaviour. The semi-natural experiments confirmed these results as interspecific competition affected relationships between individual growth rate of salmon, initial weight and activity index. Overall, individual-based analyses highlighted important mechanisms that were concealed at the group level, and that may be crucial to understand ecological and evolutionary consequences of exotic species. Moreover, these results demonstrated that competition with an exotic species disrupts the hierarchical relationship among native individuals and may therefore represent a potential for a shift in selective pressure.     

The diversity of juvenile salmonids does not affect their competitive impact on a native galaxiid

We used an invaded stream fish community in southern Chile to experimentally test whether the diversity of exotic species affects their competitive impact on a native species. In artificial enclosures an established invasive, rainbow trout, Oncorhynchus mykiss, and a potential invader, Atlantic salmon, Salmo salar, reduced the growth rate of native peladilla, Aplochiton zebra, by the same amount. In enclosures with both exotic salmonids, the growth rates of all three species were the same as in single exotic treatments. While neither species identity nor diversity appeared to affect competitive interactions in this experiment, the impact of salmonid diversity may vary with the type of interspecific interaction and/or the species identity of the exotics. Our experiment links two prominent concepts in invasion biology by testing whether the result of invasional meltdown, an increase in the diversity of exotic species, affects their impact through interspecific competition, the mechanism invoked by the biotic resistance hypothesis.     

Comparison of competitive ability between native and introduced salmonids: evidence from pairwise contests

Brown trout, Salmo trutta, and rainbow trout, Oncorhynchus mykiss, have been introduced to freshwaters in Hokkaido, Japan. Today, it is recognized that these introduced salmonids have negative impacts on native salmonids such as white-spotted charr, Salvelinus leucomaenis, and masu salmon, O. masou. In particular, interspecific competition may be an important mechanism that could contribute to the exclusion for native salmonids. In this study, experimental pairwise contests were conducted to compare interference competitive ability between native and introduced salmonids. We demonstrated that brown trout were competitively superior to white-spotted charr and masu salmon whereas rainbow trout were superior to white-spotted charr. We suggest that introduced brown trout negatively impact both white-spotted charr and masu salmon, and introduced rainbow trout negatively impact white-spotted charr.     

Varying effects of anadromous sockeye salmon on the trophic ecology of two species of resident salmonids in southwest Alaska

1. Anadromous salmon transport marine‐derived nutrients and carbon to freshwater and riparian ecosystems upon their return to natal spawning systems. The ecological implications of these subsidies on the trophic ecology of resident fish remain poorly understood despite broad recognition of their potential importance. 2. We studied the within‐year changes in the ration size, composition and stable isotope signature of the diets of two resident salmonids (rainbow trout, Oncorhynchus mykiss; Arctic grayling, Thymallus arcticus) before and after the arrival of sockeye salmon (Oncorhynchus nerka) to their spawning grounds in the Bristol Bay region of southwest Alaska. 3. Ration size and energy intake increased by 480–620% for both species after salmon arrived. However, the cause of the increases differed between species such that rainbow trout switched to consuming salmon eggs, salmon flesh and blowflies that colonized salmon carcasses, whereas grayling primarily ate more benthic invertebrates that were presumably made available because of physical disturbances by spawning salmon. 4. We also observed an increase in the δ¹⁵N of rainbow trout diets post‐salmon, but not for grayling. This presumably led to the observed increase in the δ¹⁵N of rainbow trout with increasing body mass, but not for grayling. 5. Using a bioenergetics model, we predicted that salmon‐derived resources contributed a large majority of the energy necessary for growth in this resident fish community. Furthermore, the bioenergetics model also showed how seasonal changes in diet affected the stable isotope ratios of both species. These results expand upon a growing body of literature that highlights the different pathways whereby anadromous salmon influence coastal ecosystems, particularly resident fish.     

Effects of aquaculture related stressors and nutritional restriction on circulating growth factors (GH, IGF-I and IGF-II) in Atlantic salmon and rainbow trout

The effects of aquaculture related stressors on circulating levels of GH, IGF-I and for the first time, IGF-II in Atlantic salmon (Salmo salar) and rainbow trout (Oncorhynchus mykiss) were investigated. Specifically, circulating growth factor levels were measured in four different experiments. Two 24 h confinement stressor procedures, (one with Atlantic salmon, the other with rainbow trout); following a hypo-osmotic stressor (freshwater bath) in salt water acclimated, adult, Atlantic salmon; and during a 22 day starvation and re-feeding protocol with juvenile Atlantic salmon. Handling and confinement resulted in significant decreases in circulating levels of all three growth factors in Atlantic salmon, and IGF-I and IGF-II (but not GH) in rainbow trout. A 2-3 h freshwater bath to remove gill parasites on a commercial Atlantic salmon aquaculture operation caused a significant decrease in circulating GH and IGF-I concentrations, but no significant change in IGF-II concentration, 2 days post bathing. Starvation for a period of 15 days in Atlantic salmon resulted in a significant increase in circulating GH levels and a significant decrease in circulating IGF-I and IGF-II. Re-feeding of starved fish for 7 days resulted in a significant decrease in GH to the concentration measured in continually fed fish, however re-feeding did not change plasma levels of IGF-I and IGF-II relative to continually starved fish. The results presented here confirm previously observed handling and confinement stressor induced effects on GH and IGF-I and, for the first time, on IGF-II in salmonids. Furthermore this study confirms the nutritional regulation of GH, IGF-I and IGF-II in juvenile Atlantic salmon.     

Skin morphology and humoral non-specific defence parameters of mucus and plasma in rainbow trout,coho and Atlantic salmon

Susceptibility to different diseases among related species, such as coho salmon (Oncorhynchus kisutch), rainbow trout (Oncorhyncus mykiss) and Atlantic salmon (Salmo salar), is variable. The prominence of these species in aquaculture warrants investigation into sources of this variability to assist future disease management. To develop a better understanding of the basis for species variability, several important non-specific humoral parameters were examined in juvenile fish of these three economically important species. Mucous protease, alkaline phosphatase and lysozyme, as well as plasma lysozyme activities and histological parameters (epidermal thickness and mucous cell density, and size) were characterized and compared for three salmonids: rainbow trout, Atlantic salmon and coho salmon. Rainbow trout had a thicker epidermis and significantly more mucous cells per cross-sectional area than the other two species. Rainbow trout also had significantly higher mucous protease activity than Atlantic salmon and significantly higher lysozyme (plasma and mucus) activities than coho and Atlantic salmon, in seawater. Atlantic salmon, on the other hand, had the lowest activities of mucous lysozyme and proteases, the thinnest epidermal layer and the sparsest distribution of mucous cells, compared with the two other salmonids in seawater. Only coho salmon had sacciform cells. Atlantic and coho salmon had higher mucous lysozyme activities in freshwater as compared to seawater. There was no significant difference between mucous lysozyme activities in any of the three species reared in freshwater; however, rainbow trout still had a significantly higher plasma lysozyme activity compared with the other two species. All three species exhibited significantly lower mucous alkaline phosphatase and protease activities in freshwater than in seawater. Our results demonstrate that there are significant histological and biochemical differences between the skin and mucus of these three salmonid species, which may change as a result of differing environments. Variation in these innate immune factors is likely to have differing influences on each species response to disease processes.     

The viscosity and glycoprotein biochemistry of salmonid mucus varies with species,salinity and the presence of amoebic gill disease

Fish mucus has previously been reported to change in appearance and composition among species and in response to changes in salinity and disease status. This study reports on the mucus viscosity and glycoprotein biochemistry of Atlantic salmon (Salmo salar L.), brown trout (Salmo trutta L.) and rainbow trout (Oncorhynchus mykiss Walbaum) in freshwater and seawater, both naïve to and affected by amoebic gill disease (AGD). Cutaneous mucus viscosity was measured over a range of shear rates (11.5, 23, 46 and 115 s^–1), and non-Newtonian behaviour was demonstrated for all three species. Mucus viscosity was significantly greater in seawater than in freshwater for all species, and significantly lower in AGD-affected Atlantic salmon and brown trout. Mucus glucose, total protein and osmolality data indicated that differences in viscosity due to salinity were mostly attributed to changes in mucus hydration, while differences due to disease were mostly attributed to changes in mucus composition. Trends in gill mucus cell histochemistry included shifts in glycoproteins from neutral mucins in freshwater to acidic mucins in seawater, and shifts towards neutral mucins, with an increase in mucus cell numbers, in response to AGD. Results suggested that Atlantic salmon and brown trout are more similar to one another in their mucus profile than to rainbow trout. Atlantic salmon and brown trout both exhibited a whole-body mucus response to AGD, whereas rainbow trout exhibited only a local gill response. Findings hold implications for fish physiology and pathology, and indicate that future fish-disease management strategies should be species and condition specific.Communicated by I.D. HumeThe word mucus has been used in its noun form throughout the paper for clarity

---

An erratum to this article can be found at .     

Seasonality and heterogeneity of live fish movements in Scottish fish farms

Movement of live animals is a key contributor to disease spread. Farmed Atlantic salmon Salmo salar, rainbow trout Onchorynchus mykiss and brown/sea trout Salmo trutta are initially raised in freshwater (FW) farms; all the salmon and some of the trout are subsequently moved to seawater (SW) farms. Frequently, fish are moved between farms during their FW stage and sometimes during their SW stage. Seasonality and differences in contact patterns across production phases have been shown to influence the course of an epidemic in livestock; however, these parameters have not been included in previous network models studying disease transmission in salmonids. In Scotland, farmers are required to register fish movements onto and off their farms; these records were used in the present study to investigate seasonality and heterogeneity of movements for each production phase separately for farmed salmon, rainbow trout and brown/sea trout. Salmon FW-FW and FW-SW movements showed a higher degree of heterogeneity in number of contacts and different seasonal patterns compared with SW-SW movements. FW-FW movements peaked from May to July and FW-SW movements peaked from March to April and from October to November. Salmon SW-SW movements occurred more consistently over the year and showed fewer connections and number of repeated connections between farms. Therefore, the salmon SW-SW network might be treated as homogeneous regarding the number of connections between farms and without seasonality. However, seasonality and production phase should be included in simulation models concerning FW-FW and FW-SW movements specifically. The number of rainbow trout FW-FW and brown/sea trout FW-FW movements were different from random. However, movements from other production phases were too low to discern a seasonal pattern or differences in contact pattern.     

A global assessment of salmon aquaculture impacts on wild salmonids

Since the late 1980s, wild salmon catch and abundance have declined dramatically in the North Atlantic and in much of the northeastern Pacific south of Alaska. In these areas, there has been a concomitant increase in the production of farmed salmon. Previous studies have shown negative impacts on wild salmonids, but these results have been difficult to translate into predictions of change in wild population survival and abundance. We compared marine survival of salmonids in areas with salmon farming to adjacent areas without farms in Scotland, Ireland, Atlantic Canada, and Pacific Canada to estimate changes in marine survival concurrent with the growth of salmon aquaculture. Through a meta-analysis of existing data, we show a reduction in survival or abundance of Atlantic salmon; sea trout; and pink, chum, and coho salmon in association with increased production of farmed salmon. In many cases, these reductions in survival or abundance are greater than 50%. Meta-analytic estimates of the mean effect are significant and negative, suggesting that salmon farming has reduced survival of wild salmon and trout in many populations and countries.     

Emergence and maintenance of infectious salmon anaemia virus (ISAV) in Europe: a new hypothesis

The present study describes the use of molecular methods in studying infectious salmon anaemia virus (ISAV), an important pathogen of farmed salmon in Norway, Scotland, the Faeroe Islands, Canada, USA and Chile. The nucleotide sequences of the haemagglutinin gene (HA) from 70 ISAV isolates have been analysed for phylogenetic relationship and the average mutation rate of nucleotide substitutions calculated. The isolates constitute 2 major groups, 1 European and 1 North American group. The isolate from Chile is closely related to the North American isolates. The European isolates can be further divided into 3 separate groups reflecting geographical distribution, time of collection, and transmission connected with farming activity. Based on existing information about infectious salmon anaemia (ISA) and new information emerging from the present study, it is hypothesised that: (1) ISAV is maintained in wild populations of trout and salmon in Europe; (2) it is transmitted between wild hosts mainly during their freshwater spawning phase in rivers; (3) wild salmonids, mainly trout, possibly carry benign wild-type ISAV isolates; (4) a change (mutation) in virulence probably results from deletions of amino acid segments from the highly polymorphic region (HPR) of benign wild-type isolates; (5) ISA emerges in farmed Atlantic salmon when mutated isolates are transmitted from wild salmonids or, following mutation of benign isolates, in farmed salmon after transmission from wild salmonids; (6) farming activity is an important factor in transmission of ISAV between farming sites in addition to transmission of ISAV from wild salmonids to farmed salmon; (7) transmission of ISAV from farmed to wild salmonids probably occurs less frequently than transmission from wild to farmed fish due to lower frequency of susceptible wild individuals; (8) the frequency of new outbreaks of ISA in farmed salmon probably reflects natural variation in the prevalence of ISAV in wild populations of salmonids.     

Serum agglutinating titres against Renibacterium salmoninarum the causative agent of bacterial kidney disease, in rainbow trout, Salmo gairdneri Richardson, and Atlantic salmon, Salmo salar L.

Serum agglutinating titres against Renibacterium salmoninarum were determined from clinically infected, sub-clinically infected and non-infected rainbow trout, Salmo gairdneri Richardson, and Atlantic salmon, Salmo salar L. Titres ≥16 occurred in clinically infected rainbow trout, but declined after a few months when disease signs were absent. Non-infected rainbow trout had titres ≤ 16. No correlation between the agglutinating titre and the level of infection was found in farmed Atlantic salmon. Variable titres were found in wild salmon broodstock although there was no evidence of BKD during 4 years of testing.     

Predictability of multispecies competitive interactions in three populations of Atlantic salmon Salmo salar

Juvenile Atlantic salmon Salmo salar from three allopatric populations (LaHave, Sebago and Saint‐Jean) were placed into artificial streams with combinations of four non‐native salmonids: brown trout Salmo trutta, rainbow trout Oncorhynchus mykiss, Chinook salmon Oncorhynchus tshawytscha and coho salmon Oncorhynchus kisutch. Non‐additive effects, as evidenced by lower performance than predicted from weighted summed two‐species competition trials, were detected for S. salar fork length (L_F) and mass, but not for survival, condition factor or riffle use. These data support emerging theory on niche overlap and species richness as factors that can lead to non‐additive competition effects.     

Competition,predation and flow rate as mediators of direct and indirect effects in a stream food chain

Using semi-natural stream channels, we estimated the effects of competition and predation exerted by juvenile and adult exotic rainbow trout (Oncorhynchus mykiss) on the diel activity pattern of juvenile native Atlantic salmon (Salmo salar), a secondary consumer. We also evaluated the direct and indirect effects of competition, predation and abiotic factors (water depth and velocity) on the growth rate of salmon, the biomass of invertebrate grazers (primary consumers) and the biomass of periphytic algae (primary producers; chlorophyll a). The presence of chemical cues emanating from adult predatory trout reduced the daily activity of juvenile Atlantic salmon. In contrast, competition imposed by juvenile rainbow trout forced Atlantic salmon to be more active during the day, even if adult rainbow trout were also present. We found no effect of either competition or of predatory cues on the growth rate of Atlantic salmon, and no evidence of indirect effects on either the biomass of invertebrates or the biomass of chlorophyll a. In contrast, we demonstrated that this food chain (fish--invertebrate grazers--periphytic algae) was under the control of a critical abiotic factor, the water velocity, and of bottom-up processes. We concluded that the exotic species directly increases the risk of predation of the native Atlantic salmon, but behavioral compensation probably limits the effects on growth rate. The competition and predation imposed by the invaders had no indirect effects on lower trophic levels. Top-down effects may have been mitigated by the dominant influence of water velocity controlling all components of the food chain and by elevated levels of primary production.     

Previously unrecognised division within Moritella viscosa isolated from fish farmed in the North Atlantic

Previously undocumented phenotypical and genetic variation was identified amongst isolates of Moritella viscosa collected from various geographical locations and from different fish species. The studied isolates could be split into 2 major phenotypically and genetically different clusters, one of which was consistent with the species type strain (NCIMB 13548). Isolates consistent with the type strain originated exclusively from Atlantic salmon farmed in Norway, Scotland and the Faroe Isles, although a single isolate from farmed Norwegian cod clustered closely with this group. The 'variant' cluster comprised isolates originating from Norwegian farmed rainbow trout, Icelandic farmed rainbow trout and salmon, Canadian farmed (Atlantic) salmon, Icelandic lumpsucker and only exceptionally from Norwegian salmon. With the exception of the single aforementioned cod isolate, all isolates from Norwegian farmed cod belonged to the variant cluster. Phenotypically, the clusters could be absolutely separated only by elevated haemolytic activity in the variant strain, although approximately half of these isolates also produced acid from mannose, in contrast to the typical (type) strain. While 16S rRNA gene sequencing was unable to separate the 2 clusters, Western blot analyses, plasmid profile analysis, pulsed field gel electrophoresis and gyrB gene sequence analysis produced clusters consistent with the phenotypic data. Macroscopically and histologically the disease in rainbow trout caused by the variant strain was consistent with that previously described in Atlantic salmon. The results of the present study may indicate a degree of host specificity of the typical strain for Atlantic salmon.     

Prevalence and diagnosis of bacterial kidney disease (BKD) in Scotland between 1990 and 2002

Bacterial kidney disease (BKD) is a notifiable disease for salmonids under United Kingdom and European Union legislation. Within the UK, legislation and the control of infected fish with BKD has been operating for 25 yr. Infection by the bacterium Renibacterium salmoninarum results in a chronic, debilitating infection and mortality. Records of BKD outbreaks and the detection of R. salmoninarum were monitored from 1990 through to 2002 for Atlantic salmon Salmo salar and rainbow trout Oncorhynchus mykiss reared in Scottish waters. The test methods included ELISA, culture and light microscopy. New outbreaks of BKD in salmon in seawater declined during this period, but with year-to-year variation. Only 1 record of BKD has occurred in freshwater-reared salmon (prevalence 1.04). BKD in farmed rainbow trout in seawater is uncommon and was only identified in 1993 and between 1998 and 2000. The number of active designated area orders (DAOs) for outbreaks in salmon has fallen since 1990, but has remained relatively constant for trout over the period of study.     

Prevalence of infectious salmon anaemia virus (ISAV) in wild salmonids in western Norway

Studies of infectious salmon anaemia virus (ISAV), an important pathogen of farmed salmon in Norway, Scotland, the Faeroe Islands, Ireland, Canada, the USA and Chile, suggest that natural reservoirs for this virus can be found on both sides of the North Atlantic. Based on existing information about ISAV it is believed to be maintained in wild populations of trout and salmon in Europe. It has further been suggested that ISAV is transmitted between wild hosts, mainly during their freshwater spawning phase in rivers, and that wild salmonids, mainly trout, are possible carriers of benign wild-type variants of ISAV. Change in virulence is probably a result of deletions of amino acid segments from the highly polymorphic region (HPR) of benign wild-type isolates after transmission to farmed salmon. Hence, it has been suggested that the frequency of new outbreaks of ISA in farmed salmon could partly reflect natural variation in the prevalence of ISAV in wild populations of salmonids. The aims of the present study were to screen for ISAV in wild salmonids during spawning in rivers and to determine the pathogenicity of resultant isolates from wild fish. Tissues from wild salmonids were screened by RT-PCR and real-time PCR. The prevalence of ISAV in wild trout Salmo trutta varied from 62 to 100% between tested rivers in 2001. The prevalence dropped in 2002, ranging from 13 to 36% in the same rivers and to only 6% in 2003. All ISAV were nonpathogenic when injected into disease-free Atlantic salmon, but were capable of propagation, as indicated by subsequent viral recovery. However, non-pathogenic ISAV has also been found in farmed salmon, where a prevalence as high as 60% has been registered, but with no mortalities occurring. Based on the results of the present and other studies, it must be concluded that vital information about the importance of wild and man-made reservoirs for the emergence of ISA in salmon farming is still lacking. This information can only be gained by further screening of possible reservoirs, combined with the development of a molecular tool for typing virulence and the geographical origin of the virus isolates.     

Genetic differentiation of populations of the copepod sea louse Lepeophtheirus salmonis (Krøyer) ectoparasitic on wild and farmed salmonids around the coasts of Scotland: Evidence from RAPD markers

Sea trout are the sea-going migratory form of the freshwater brown trout (Salmo trutta L.) and since 1989 there have been marked declines in their stocks on the west coasts both of Scotland and Ireland. Various factors have been attributed as possible causal agents in these stock declines, including fresh water acidification, overfishing, climatic fluctuations, habitat degradation and sea lice parasitic burdens. The putative impact of infestations of sea trout by the ectoparasitic copepod sea louse, Lepeophtheirus salmonis (Krøyer), has featured prominently in the controversy, especially with regard to the role of inshore commercial salmon farms as a possible source of infestation of wild salmonids by sea lice. This study focused on the population genetics of L. salmonis around the coasts of Scotland: We sampled fish from wild and cultured stocks and included salmon (Salmo salar L.), rainbow trout (Oncorhynchus mykiss Walbaum) and sea trout as host species. Analyses of allozyme variation of sea lice were confined to data for two polymorphic loci (Fum, Got-2) and conformed to our initial expectation — that the inclusion of a planktonic larval phase in the life cycle of the copepod, in addition to the high mobility of the host fish, would enhance gene flow and preclude genetic differentiation of L. salmonis populations as a result of random drift alone. DNA polymorphism was quantified by means of PCR and RAPD analysis. Six primers were screened for 16 samples (from wild and farmed salmon, wild sea trout and farmed rainbow trout) — including the east, north and west coasts of Scotland — and the data analyzed by AMOVA (Analysis of Molecular Variance). In contrast to the allozyme results, the RAPD analysis showed striking patterns of genetic differentiation around the coasts of Scotland. The overall pattern was one of genetic homogeneity of L. salmonis populations sampled from wild salmon and sea trout. All of the L. salmonis samples taken from farmed salmon and rainbow trout did, however, show highly significant levels of genetic differentiation, both between wild and farmed salmonids and among the various farms themselves. Evidence of high levels of small-scale spatial or temporal heterogeneity of RAPD marker band frequencies was shown for the one farm from which repeat samples (July and November, 1995) were analysed. Samples of sea lice taken from west coast wild sea trout subjected to RAPD analysis also revealed the occurrence of putative “farm markers” in some individual parasites, indicating that they had possibly originated from salmon farms.     

Size-structured Interactions between Native and Introduced Species: Can Intraguild Predation Facilitate Invasion by Stream Salmonids?

Dynamics of biological invasions may be complicated in size-structured animal populations. Differences in timing of life history events such as juvenile emergence create complex interaction webs where different life stages of native and non-native species act as predators, competitors, and prey. Stream salmonids are an ideal group for studying these phenomena because they display competition and predation in size-structured populations and have been introduced worldwide. For example, introduced rainbow trout (Oncorhynchus mykiss) are invading streams of Hokkaido Island, Japan and have caused declines in native masu salmon (O. masou) populations. However, age-0 rainbow trout emerge later than age-0 masu salmon and are smaller, which raises the question of why they are able to recruit and therefore invade in the face of a larger competitor. We conducted experiments in laboratory stream channels to test effects of increasing density of age-0 and age-1 rainbow trout on age-0 masu salmon. Age-1 rainbow trout dominated age-0 masu salmon by aggressive interference, relegating them to less favorable foraging positions downstream and reducing their foraging frequency and growth. The age-1 trout also reduced masu salmon survival by predation of about 40% of the individuals overall. In contrast, age-0 rainbow trout had little effect on age-0 masu salmon. Instead, the salmon dominated the age-0 trout by interference competition and reduced their survival by predation of 60% of the individuals. In each case, biotic interactions by the larger species on the smaller were strongly negative due to a combination of interspecific competition and intraguild predation. We predict that together these produce a positive indirect effect in the interaction chain that will allow the recruitment of rainbow trout in the face of competition and predation from age-0 masu salmon, and thereby facilitate their invasion in northern Japan.     

Sustainability of the Lake Superior Fish Community: Interactions in a Food Web Context

The restoration and rehabilitation of the native fish communities is a long-term goal for the Laurentian Great Lakes. In Lake Superior, the ongoing restoration of the native lake trout populations is now regarded as one of the major success stories in fisheries management. However, populations of the deepwater morphotype (siscowet lake trout) have increased much more substantially than those of the nearshore morphotype (lean lake trout), and the ecosystem now contains an assemblage of exotic species such as sea lamprey, rainbow smelt, and Pacific salmon (chinook, coho, and steelhead). Those species play an important role in defining the constraints and opportunities for ecosystem management. We combined an equilibrium mass balance model (Ecopath) with a dynamic food web model (Ecosim) to evaluate the ecological consequences of future alternative management strategies and the interaction of two different sets of life history characteristics for fishes at the top of the food web. Relatively rapid turnover rates occur among the exotic forage fish, rainbow smelt, and its primary predators, exotic Pacific salmonids. Slower turnover rates occur among the native lake trout and burbot and their primary prey—lake herring, smelt, deepwater cisco, and sculpins. The abundance of forage fish is a key constraint for all salmonids in Lake Superior. Smelt and Mysis play a prominent role in sustaining the current trophic structure. Competition between the native lake trout and the exotic salmonids is asymmetric. Reductions in the salmon population yield only a modest benefit for the stocks of lake trout, whereas increased fishing of lake trout produces substantial potential increases in the yields of Pacific salmon to recreational fisheries. The deepwater or siscowet morphotype of lake trout has become very abundant. Although it plays a major role in the structure of the food web it offers little potential for the restoration of a valuable commercial or recreational fishery. Even if a combination of strong management actions is implemented, the populations of lean (nearshore) lake trout cannot be restored to pre-fishery and pre-lamprey levels. Thus, management strategy must accept the ecological constraints due in part to the presence of exotics and choose alternatives that sustain public interest in the resources while continuing the gradual progress toward restoration. Received 10 December 1999; accepted 13 June 2000.