Reproductive Success of Captive-Bred Steelhead Trout in the Wild: Evaluation of Three Hatchery Programs in the Hood River

Abstract:  Population supplementation programs that release captive-bred offspring into the wild to boost the size of endangered populations are now in place for many species. The use of hatcheries for supplementing salmonid populations has become particularly popular. Nevertheless, whether such programs actually increase the size of wild populations remains unclear, and predictions that supplementation fish drag down the fitness of wild fish remain untested. To address these issues, we performed DNA-based parentage analyses on almost complete samples of anadromous steelhead ( Oncorhynchus mykiss ) in the Hood River in Oregon (U.S.A.). Steelhead from a supplementation hatchery (reared in a supplementation hatchery and then allowed to spawn naturally in the wild) had reproductive success indistinguishable from that of wild fish. In contrast, fish from a traditional hatchery (nonlocal origin, multiple generations in hatcheries) breeding in the same river showed significantly lower fitness than wild fish. In addition, crosses between wild fish and supplementation fish were as reproductively successful as those between wild parents. Thus, there was no sign that supplementation fish drag down the fitness of wild fish by breeding with them for a single generation. On the other hand, crosses between hatchery fish of either type (traditional or supplementation) were less fit than expected, suggesting a possible interaction effect. These are the first data to show that a supplementation program with native brood stock can provide a single-generation boost to the size of a natural steelhead population without obvious short-term fitness costs. The long-term effects of population supplementation remain untested.     

Use of Linkage Disequilibrium Data to Estimate Effective Size of Hatchery and Natural Fish Populations

A primary parameter in the assessment of the viability of a population is its effective population size ( Ne ). Allozyme analysis of four groups of fishes provided data on linkage disequilibrium, which were then used to estimate Ne . The groups included hatchery samples of juvenile white seabass, Atractoscion nobilis , juvenile rainbow trout, Oncorhynchus mykiss , from the Shasta Hatchery, and juvenile chinook salmon, O. tshawytscha , from the Coleman National Fish Hatchery. The fourth sample consisted of juvenile chinook salmon from the threatened winter run in the upper Sacramento River. The groups of fish were chosen to represent different applications of the methodology to conservation of fishes. For a variety of reasons. Ne may be considerably lower than census counts of fish present in the parental populations. The Ne of the hatchery broodstock that produced the sample of juvenile white seabass was estimated to be approximately 10, although 25 adult white seabass were present in a mass spawning tank. Ne estimates for the parental populations of the Shasta and Coleman Hatchery samples were 35.8 and 132.5, respectively. The actual number of fish spawned at the Shasta Hatchery was approximately 40, whereas nearly 10,000 salmon were spawned at the Coleman Hatchery. The threatened winter run of chinook salmon had an estimated Ne of 85.5 and an approximate run size of 2000 salmon. The method of estimating effective population size from linkage disequilibrium data appears to result in realistic estimates of effective population size when adequate sample size and a sufficient number of polymorphic loci are available.     

Human Influence on the Spatial Structure of Threatened Pacific Salmon Metapopulations

Abstract: To remain viable, populations must be resilient to both natural and human‐caused environmental changes. We evaluated anthropogenic effects on spatial connections among populations of Chinook salmon (Oncorhynchus tshawytscha) and steelhead (O. mykiss) (designated as threatened under the U.S. Endangered Species Act) in the lower Columbia and Willamette rivers. For several anthropogenic‐effects scenarios, we used graph theory to characterize the spatial relation among populations. We plotted variance in population size against connectivity among populations. In our scenarios, reduced habitat quality decreased the size of populations and hydropower dams on rivers led to the extirpation of several populations, both of which decreased connectivity. Operation of fish hatcheries increased connectivity among populations and led to patchy or panmictic populations. On the basis of our results, we believe recolonization of the upper Cowlitz River by fall and spring Chinook and winter steelhead would best restore metapopulation structure to near‐historical conditions. Extant populations that would best conserve connectivity would be those inhabiting the Molalla (spring Chinook), lower Cowlitz, or Clackamas (fall Chinook) rivers and the south Santiam (winter steelhead) and north fork Lewis rivers (summer steelhead). Populations in these rivers were putative sources; however, they were not always the most abundant or centrally located populations. This result would not have been obvious if we had not considered relations among populations in a metapopulation context. Our results suggest that dispersal rate strongly controls interactions among the populations that comprise salmon metapopulations. Thus, monitoring efforts could lead to understanding of the true rates at which wild and hatchery fish disperse. Our application of graph theory allowed us to visualize how metapopulation structure might respond to human activity. The method could be easily extended to evaluations of anthropogenic effects on other stream‐dwelling populations and communities and could help prioritize among competing conservation measures.     

Does juvenile experience affect migration and spawning of adult Atlantic salmon?

Summary This paper documents differences in seasonal time of river ascent and descent, and instream behavior of adult wild and sea-ranched Atlantic salmon (Salmo salar) of the Norwegian River Imsa stock during the period 1981–1989. Wild fish use River Imsa as a nursery, and at an age of 2 years most of them migrate to the sea as smolts. The sea ranched fish are hatchery reared offspring of the River Imsa stock and are released as smolts at the mouth of the river. They are thus deprived of juvenile river life and a downstream smolt migration. Wild and sea ranched salmon feed for 1 or more years in the Norwegian Sea before homing as spawners. Both groups returned simultaneously to coastal Norway, but sea ranched fish ascended the river later and descended sooner after spawning than wild fish. All wild females and almost all wild males (96.2%) spawned in the river, whereas 13.5% and 36.7%, respectively, of the mature sea-ranched females and males left the river unspawned. The annual number, but not the proportion, of unspawned fish increased with increasing density of adult salmon in the river. Unspawned females were medium sized and small (45–70 cm); unspawned males were medium sized and large (50–90 cm). Independent of the density of spawners in the river, sea ranched fish moved up- and downstream the river more often than wild fish. More than 20% of the sea-ranched salmon and less than 1% of the wild salmon passed a trap 100 m above the river outlet more than once in each direction during the same spawning reason. Moreover, sea-ranched salmon were about twice as often seriously injured during spawning as wild fish. Lack of juvenile experience from the river may be the main reason for the behavioral differences between sea-ranched and wild fish. Offprint requests to: B. Jonsson     

Naturally and hatchery produced European trout Salmo trutta: do their marine survival and dispersal differ?

We tested whether marine survival and migration pattern differed between naturally and hatchery produced European trout Salmo trutta of different origins. The hatchery fish were released 150 m above the river estuary of the southwestern, Norwegian River Imsa, the home of the local population. Recaptures were used as proxy for survival. Wild and local hatchery fish survived better than transplanted hatchery stocks. Trout that were 1 year at release survived less well than 2-year olds, and small individuals less well than larger ones. Relative to their body size at release, populations that originated most distant from the River Imsa, the Baltic River Emån and the Norwegian mountain Lake Tunhovd, exhibited the poorest sea survival. At sea, trout chiefly moved less than 240 km from the river of release, but there were significant differences in dispersal among populations. Hatchery-produced River Emån and Lake Tunhovd trout moved farther from the River Imsa than the south Norwegian sea trout populations, and the marine distributions of the former were similar to that of the natural River Imsa trout. Large fish moved farther from the river than smaller ones. Straying to other rivers was low among wild and local hatchery-produced fish, and significantly lower than among most transplanted populations, and River Emån trout in particular. Thus, the River Imsa trout appeared better adapted to survival under the local conditions than non-local trout with consequences for optimal population management.     

Evaluating Alternative Strategies for Minimizing Unintended Fitness Consequences of Cultured Individuals on Wild Populations

Artificial propagation strategies often incur selection in captivity that leads to traits that are maladaptive in the wild. For propagation programs focused on production rather than demographic contribution to wild populations, effects on wild populations can occur through unintentional escapement or the need to release individuals into natural environments for part of their life cycle. In this case, 2 alternative management strategies might reduce unintended fitness consequences on natural populations: (1) reduce selection in captivity as much as possible to reduce fitness load (keep them similar), or (2) breed a separate population to reduce captive‐wild interactions as much as possible (make them different). We quantitatively evaluate these 2 strategies with a coupled demographic–genetic model based on Pacific salmon hatcheries that incorporates a variety of relevant processes and dynamics: selection in the hatchery relative to the wild, assortative mating based on the trait under selection, and different life cycle arrangements in terms of hatchery release, density dependence, natural selection, and reproduction. Model results indicate that, if natural selection only occurs between reproduction and captive release, the similar strategy performs better. However, if natural selection occurs between captive release and reproduction, the different and similar strategies present viable alternatives to reducing unintended fitness consequences because of the greater opportunity to purge maladaptive individuals. In this case, the appropriate approach depends on the feasibility of each strategy and the demographic goal (e.g., increasing natural abundance, or ensuring that a high proportion of natural spawners are naturally produced). In addition, the fitness effects of hatchery release are much greater if hatchery release occurs before (vs. after) density‐dependent interactions. Given the logistical challenges to achieving both the similar and different strategies, evaluation of not just the preferred strategy but also the consequences of failing to achieve the desired target is critical. Evaluación de Estrategias Alternativas para Minimizar las Consecuencias No Inesperadas en la Adecuación de Individuos Criados en Cautiverio sobre Poblaciones Silvestres     

Effective Population Size in Winter-Run Chinook Salmon

Winter-run chinook salmon from the Sacramento River, California, is federally listed as endangered. Since 1989 there has been aprogram to augment the natural population by capturing adults, artificially spawning them, raising tine young and releasing the smolt. Here we estimate the effective population size of these captive-raised fish, the natural run, and the combination of both groups over the three-year period from 1991 to 1993. We find that the most appropriate estimate of the effective population size of the captive-raised progeny is a variance estimate of effective population size standardized so that the number of released smolts returning to spawn was the same as the number of spawners used to produce the smolts originally. We have generated 10,000 random samples to simulate returns from these released progeny. The estimates of the effective population sizes in 1991, 1992, and 1993 were only 7.02, 19.07, and 7.74, respectively. We then determined limits on the effective population size of the natural run based on 0.1 and 0.333 of the run-size estimates. Using estimates of the captive proportion of the run, the minimum estimates of the effective population size of the overall run for the three years were 21.9, 127.3, and 39.0, and the maximum estimates were 61.6, 401.0, and 108.7. It does not appear that the hatchery program has reduced the overall effective population size. The run sizes in each year are extremely low, however, and it is possible that fish will be lost from this run in one of the years in the immediate future, making reestablishment of a healthy run even more difficult.     

Interacting Effects of Translocation,Artificial Propagation,and Environmental Conditions on the Marine Survival of Chinook Salmon from the Columbia River,Washington, U.S.A.

Abstract: Captive rearing and translocation are often used concurrently for species conservation, yet the effects of these practices can interact and lead to unintended outcomes that may undermine species’ recovery efforts. Controls in translocation or artificial‐propagation programs are uncommon; thus, there have been few studies on the interacting effects of these actions and environmental conditions on survival. The Columbia River basin, which drains 668,000 km² of the western United States and Canada, has an extensive network of hydroelectric and other dams, which impede and slow migration of anadromous Pacific salmon (Oncorhynchus spp.) and can increase mortality rates. To mitigate for hydrosystem‐induced mortality during juvenile downriver migration, tens of millions of hatchery fish are released each year and a subset of wild‐ and hatchery‐origin juveniles are translocated downstream beyond the hydropower system. We considered how the results of these practices interact with marine environmental conditions to affect the marine survival of Chinook salmon (O. tshawytscha). We analyzed data from more than 1 million individually tagged fish from 1998 through 2006 to evaluate the probability of an individual fish returning as an adult relative to its rearing (hatchery vs. wild) and translocation histories (translocated vs. in‐river migrating fish that traveled downriver through the hydropower system) and a suite of environmental variables. Except during select periods of very low river flow, marine survival of wild translocated fish was approximately two‐thirds less than survival of wild in‐river migrating fish. For hatchery fish, however, survival was roughly two times higher for translocated fish than for in‐river migrants. Competition and predator aggregation negatively affected marine survival, and the magnitude of survival depended on rearing and translocation histories and biological and physical conditions encountered during their first few weeks of residence in the ocean. Our results highlight the importance of considering the interacting effects of translocation, artificial propagation, and environmental variables on the long‐term viability of species.     

Temperature-associated population diversity in salmon confers benefits to mobile consumers

Habitat heterogeneity can generate intraspecific diversity through local adaptation of populations. While it is becoming increasingly clear that population diversity can increase stability in species abundance, less is known about how population diversity can benefit consumers that can integrate across population diversity in their prey. Here we demonstrate cascading effects of thermal heterogeneity on trout-salmon interactions in streams where rainbow trout rely heavily on the seasonal availability of anadromous salmon eggs. Water temperature in an Alaskan stream varied spatially from 5 degrees C to 17.5 degrees C, and spawning sockeye salmon showed population differentiation associated with this thermal heterogeneity. Individuals that spawned early in cool regions of the 5 km long stream were genetically differentiated from those spawning in warmer regions later in the season. Sockeye salmon spawning generates a pulsed resource subsidy that supports the majority of seasonal growth in stream-dwelling rainbow trout. The spatial and temporal structuring of sockeye salmon spawn timing in our focal stream extended the duration of the pulsed subsidy compared to a thermally homogeneous stream with a single population of salmon. Further, rainbow trout adopted movement strategies that exploited the multiple pulses of egg subsidies in the thermally heterogeneous stream. Fish that moved to track the resource pulse grew at rates about 2.5 times higher than those that remained stationary or trout in the reference stream with a single seasonal pulse of eggs. Our results demonstrate that habitat heterogeneity can have important effects on the population diversity of dominant species, and in turn, influence their value to species that prey upon them. Therefore, habitat homogenization may have farther-reaching ecological effects than previously considered.     

Sexual selection on mature male parr of masu salmon (Oncorhynchus masou): does sneaking behavior favor small body size and less-developed sexual characters?

In many salmonid species, males exhibit morphological dimorphism associated with alternative mating behaviors. ”Precocious males” have a small body size with little or no development of sexual characters and adopt sneaking to gain access to females, while ”migratory males” of large body size and well-developed secondary sexual characters fight. We quantified selection on precocious male parr of masu salmon (Oncorhynchus masou) under simulated natural conditions to examine the contribution of morphology to sneaking success. In contrast to the prediction that sneaking behavior favors small body size, we detected selection favoring relatively large body size for sneaking success. This selection pressure was caused by the dominance hierarchy within parr and may have been facilitated by indifference of dominant migratory males to parr. Unlike the secondary sexual characters exhibited by migratory male salmon, such as the hooked snout and humped back, no morphological characters other than body size contributed to the reproductive success of masu salmon parr. This non-contribution may have been responsible for the lack of development of sexual characters in precocious males. Received: 15 November 1999 / Accepted: 20 May 2000     

Operational sex ratio and alternative reproductive behaviours in the European bitterling,<Emphasis Type="Italic"> Rhodeus sericeus</Emphasis>

We investigated the effects of male population density and male-biased operational sex ratio (OSR) with constant and limited resource density on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus. This species spawns inside living unionid mussels. Large males defended territories and were aggressive towards conspecifics under equal sex ratios. They also monopolised pair spawnings with females, releasing 98% of all sperm clouds during mating. However, the mating tactic changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. Large, medium and small males now obtained 61%, 33%, and 6% of sperm releases respectively, thereby reducing the opportunity for sexual selection by half. Females spawned at equal rates in the two densities of males, despite lower courtship at high density. These results run counter to the usual expectation that an increasingly male-biased OSR should lead to higher variance in male mating success. Instead, the use of alternative reproductive behaviours by males can lead to lower resource competition and mating variance at high male densities.     

Techno-Arrogance and Halfway Technologies: Salmon Hatcheries on the Pacific Coast of North America

Humankind has adopted an arrogant and ultimately self-defeating attitude toward nature that places technological mastery over nature at the forefront of our approach to many environmental problems. This "techno-arrogance" fails to recognize limitations on, and ramifications of, attempted control of nature. An example of techno-arrogance is the flawed attempt to recover Pacific salmonid fisheries through technological application in the form of hatcheries. Countless salmon stocks have declined precipitously over the last century as a result of overfishing and widespread habitat destruction. A central feature of recovery efforts has been to build many hatcheries to produce large quantities of fish to restock streams. This approach addresses the symptoms but not the causes of the declines (an example of a halfway technology), because the habitats remain largely unsuitable for salmon. There are at least six reasons why the hatchery approach will ultimately fail: (1) data demonstrate that hatcheries are not solving the problem—salmon continue to decline despite decades of hatchery production; (2) hatcheries are costly to run, and divert resources from other efforts, such as habitat restoration; (3) hatcheries are not sustainable in the long term, requiring continual input of money and energy, (4) hatcheries are a genetically unsound approach to management that can adversely affect wild populations; (5) hatchery production leads to increased harvest of declining wild populations of salmon; and (6) hatcheries conceal from the public the truth of real salmon decline. I recommend that salmonid management turn from the symptoms to the causes of decline. Overharvest and habitat destruction must be directly addressed in a major, landscape-level effort, on a scale comparable to the hatchery program, if salmonid fisheries are to remain a part of the ecological recreational, commercial and asthetic arenas in the long term.     

Investment in territorial defence depends on rearing environment in brown trout (Salmo trutta)

In many animals, territoriality will arise or cease depending on environmental factors such as intruder rate and resource availability. We investigated the effect of rearing environment on territorial behaviour in ~1.5-month-old brown trout. In the laboratory, wild-caught (reared at a low density) and hatchery-reared (high density) trout were allowed to defend a territory against a size-matched intruder reared in the same or the other environment. Because territorial behaviour should be relaxed at high-rearing densities, we hypothesized that hatchery-reared trout should value their territories less and therefore invest less in defence compared with wild-caught trout. However, in all cases, territory owners were more likely to win the contest and hatchery-reared trout were just as likely as wild-reared to win mixed contests. Furthermore, pairs of hatchery-reared trout initiated contests sooner, fought longer and were more aggressive during the contest compared with pairs of wild trout. When hatchery-reared owners met wild intruders, the contest ended sooner compared with when the roles were reversed. We conclude that territorial behaviour in brown trout is largely innate, but that the hatchery environment has promoted more aggressive individuals. These results suggest that hatchery-reared trout invest more time and energy to obtain the same contest success as wild trout. In conclusion, the lack of experience of territorial defence in a high-density rearing environment seems to reduce the efficiency of territorial behaviour. In turn, this may have negative consequences for the performance of released hatchery fish in the wild.     

Potential latitudinal variation in egg size and number of a geographically widespread reef fish,revealed by common-environment experiments

Variation in maternal reproductive traits was examined in field and reared populations of a geographically widespread reef fish, Pomacentrus coelestis (Pomacentridae), drawn from three different latitudes in Japan. Size-specific clutch size and clutch weight of wild fish increased with increasing latitude. Conversely, latitudinal variation in egg size of wild fish was obscure in same-season comparisons, probably because of the temperature effect on egg size. Common-environment experiments conducted at three temperatures showed that egg size decreased with increasing temperature in all populations. In the experiments, egg size, clutch size and clutch weight differed among populations at all temperatures, showing clear latitudinal clines. Females from low latitude spawned larger eggs at every experimental temperature. Size-specific clutch size and weight were greater in females from high latitude. Thus, the northern fish had a larger reproductive output per spawning and a larger number of smaller eggs in a spawning. Such interpopualtion variation in this fish is likely to be partially genetically based, although environmental effects on the variation cannot be entirely ruled out. This study provides evidence of potential latitudinal variation in the egg size and number in a coastal fish, by common-environment experiments. The close correspondence between latitudes and these maternal reproductive traits may be a consequence of local adaptation.     

Body size preferences in the pot-bellied seahorse <Emphasis Type="Italic">Hippocampus abdominalis:</Emphasis> choosy males and indiscriminate females

Male seahorses (genus Hippocampus) provide all post-fertilization parental care, yet despite high levels of paternal investment, these species have long been thought to have conventional sex roles, with female mate choice and male–male competition. Recent studies of the pot-bellied seahorse (Hippocampus abdominalis) have shown that sex-role reversal occurs in high-density female-biased populations, indicating that male mating preferences may lead to sexual selection on females in this species. Egg size, egg number, and offspring size all correlate positively with female body size in Hippocampus, and by choosing large mating partners, male seahorses may increase their reproductive success. While male brood size is also positively correlated with body size, small H. abdominalis males can carry exceptionally large broods, suggesting that the fecundity benefits of female preference for large partners may be limited. We investigated the importance of body size in reproductive decisions of H. abdominalis, presenting focal individuals of both sexes with potential mating partners of different sizes. Mating preferences were quantified in terms of time spent courting each potential partner. Male seahorses were highly active throughout the mate-choice trials and showed a clear behavioral preference for large partners, while females showed significantly lower levels of activity and equivocal mating preferences. The strong male preferences for large females demonstrated here suggest that sexual selection may act strongly on female body size in wild populations of H. abdominalis, consistent with predictions on the importance of female body size for reproductive output in this species. An erratum to this article can be found at     

Parental care and the cost of reproduction in a Mediterranean fish

Summary Symphodus tinca is a common near-shore Mediterranean labrid fish in which females may sometimes spawn their eggs over hundreds of square meters, or alternatively spawn into well-defined algal nests. Eggs spawned in either manner are fertilized, but widely scattered eggs receive no parental care, whereas eggs spawned into nests are usually guarded by the male until they hatch. Here, I report weight changes of individual marked fish that engaged in a variety of different reproductive behaviors during three breeding seasons. Males gained weight at 0.15% per day outside the spawning season, and added 29–78% to their overall body weight between reproductive seasons, even following substantive weight losses in a spawning season (up to 20% among nesting males). Nesting and nest-guarding males lost an average of 0.32% and 0.41% of their body weight per day in 1986 and 1987. This cost is four times greater than reproduction for nonnesting males, which registered a 0.03% daily weight gain. Actively spawning females lost 0.06% of their body weight daily during the spawning season. While long-term growth rates did not appear to be substantially affected by reproduction in either sex or by parental care in males, present work does not exclude the possibility that such long-term effects may exist.     

Exposure to variable spatial information in the early rearing environment generates asymmetries in social interactions in cod (Gadus morhua)

Many re-introduction programs used for conservation of populations and species threatened with extinction advocate the use of enriched rearing environments to train animals how to behave appropriately in the wild. Curiously, most of the current fish re-stocking programs have paid little attention to lessons previously learned in bird and mammal re-introductions. Many rehabilitation programs that use releases of hatchery fish observe higher mortality in released fish compared to wild, with most mortality arising shortly after release. One explanation for this mortality is based purely on selection processes; many hatchery fish normally selected out of the population thrive in the predator free, food-rich hatcheries. Alternatively, mortalities may be high because hatchery nursery environments fail to shape fish behaviour appropriately. Here, we empirically address the effect of enrichment in the early rearing environment in coastal cod (Gadus morhua). We find asymmetries in aggressive behaviour when fish reared in plain or enriched environments are allowed to interact. Furthermore, cod reared in standard, impoverished, hatchery environments spend less time in shelter, are more active, and show weaker anti-predator responses than fish reared with access to heterogeneous spatial cues. These results suggest that the constant, plain environments of fish farms may generate behavioural deficits that could reasonably be expected to be associated with lower survival in fish released into the wild.     

Pre- and post-mating sexual selection both favor large males in a rainbowfish

Sexual selection can act through female choice and male–male competition. Although both processes can act simultaneously, they are typically studied independently. Here, we adopt a more integrated approach to studying sexual selection by incorporating measures of both processes using the western rainbowfish Melanotaenia australis, a freshwater fish endemic to northwestern Australia. We assessed male–male competition and female choice separately while measuring the performance of individual males under both processes and used paternity analyses to estimate male reproductive success. We then related the performance of males during each of these stages to their phenotype, which was described using linear measures of size and color pattern traits, and spectrographic measures of the reflectance of color patches. We found that female choice favored relatively large males and that these preferences were consistent within individual females and repeatable between different females. Larger males were also more dominant in the competition trials and sired the majority of offspring produced when females spawned. There was little evidence to suggest that sexual selection acted on male color patterns either via female choice or male contest competition or during subsequent post-mating episodes of sexual selection. We conclude, therefore, that male–male competition and female choice act concordantly to favor relatively large males and that these patterns of mating success are reflected during post-mating episodes of sexual selection.     

Quantifying and comparing size selectivity among Alaskan sockeye salmon fisheries

Quantifying long-term size-selective harvest patterns is necessary for understanding the potential evolutionary effects on exploited species. The comparison of fishery selection patterns on the same species subject to different gear types, in different areas, and over multi-decadal periods can reveal the factors influencing selection. In this study we quantified and compared size-selective harvest by nine Alaskan sockeye salmon (Oncorhynchus nerka) fisheries to understand overall patterns. We calculated length-specific linear selection differentials (the difference in average length of fish before vs. after fishing), which are produced by different combinations of exploitation rates and length-selectivity values, and nonlinear standardized differentials, describing disruptive selection, across all years for each fishery. Selection differentials varied among years, but larger fish were caught in 73% of years for males and 84% of years for females, leaving smaller fish to spawn. Disruptive selection was observed on female and male fish in 84% and 92% of years, respectively. Linear selection was stronger on females than males in 77% of years examined, and disruptive selection was stronger on males in 71% of years. Selection pressure was influenced by a combination of factors under and beyond management control; analyses using mixed-effects models indicated that fisheries were less size selective in years when fish were larger than average and had lower exploitation rates. The observed harvest of larger than average sockeye salmon is consistent with the hypothesis that size-selective fishing contributes to decreasing age and length at maturation trends over time, but temporal variability in selection and strong disruptive selection suggests that the overall directional pressure is weaker than is often assumed in evolutionary models.     

Mating system and individual reproductive success of sympatric anadromous and resident brook charr, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>, under natural conditions

Salmonids are known for the occurrence in sympatry of two life-history forms, one that undergoes migration to sea before returning to freshwater to reproduce (anadromous) and one that inhabits freshwater without a migration phase (resident). Whereas one breeding population is often suggested by population genetic studies, mating patterns have rarely been directly assessed, especially when both sexes are found within each life-history form. By using highly polymorphic microsatellite loci and parentage analysis in a natural population of sympatric anadromous and resident brook charr (Salvelinus fontinalis), we found that gene flow occurred between the two forms and was mediated by resident males mating with both resident and anadromous females. Determinants of reproductive success, estimated by the number of surviving juveniles (ages 1 and 2 years), differed between the sexes. No strong evidence of the influence of size on individual reproductive success was found for males, whereas larger females (and hence most likely to be anadromous) were more successful. The higher individual reproductive success of anadromous fish compared to residents was mainly explained by this higher reproductive success of anadromous females. We suggest that resident males adopt a “sneaking” reproductive tactic as a way of increasing their reproductive success by mating with females of all sizes in all habitats. The persistence of the resident tactic among females may be linked to their advantage in accessing spatially constrained spawning areas in small tributary streams unavailable to larger females.