Microsatellites reveal heterosis in red deer.

The fitness consequences of inbreeding and outbreeding are poorly understood in natural populations. We explore two microsatellite-based variables, individual heterozygosity (likely to correlate with recent inbreeding) and a new individual-specific internal distance measure, mean d2 (focusing on events deeper in the pedigree), in relation to two measures of fitness expressed early in life, birth weight and neonatal survival, in 670 red deer calves (Cervus elaphus) born on the Isle of Rum between 1982 and 1996. For comparison, we also analyse inbreeding coefficients derived from pedigrees in which paternity was inferred by molecular methods. Only 14 out of 231 calves (6.1%) had non-zero inbreeding coefficients, and neither inbreeding coefficient nor individual heterozygosity was consistently related to birth weight or neonatal survival. However, mean d2 was consistently related to both fitness measures. Low mean d2 was associated with low birth weight, especially following cold Aprils, in which foetal growth is reduced. Low mean d2 was also associated with low neonatal survival, but this effect was probably mediated by birth weight because fitting birth weight to the neonatal survival model displaced mean d2 as an explanatory variable. We conclude that in the deer population fitness measures expressed early in life do not show evidence of inbreeding depression, but they do show evidence of heterosis, possibly as a result of population mixing. We also demonstrate the practical problems of estimating inbreeding via pedigrees compared with a direct marker-based estimate of individual heterozygosity. We suggest that, together, individual heterozygosity and mean d2, estimated using microsatellites, are useful tools for exploring inbreeding and outbreeding in natural population.     

Mate choice, genetic incompatibility, and outbreeding in the ornate dragon lizard, Ctenophorus ornatus

There has been recent interest in the role genetic incompatibility may play in mate or sperm choice. One source of incompatibility may be too similar or disparate genomes. An isolated population of the ornate dragon lizard, Ctenophorus ornatus, was followed over a breeding season and parentage assigned to the offspring using microsatellites. It was found that the adults in the population had an eight per cent chance of mating with a relative. I examined whether C. ornatus mate (or fertilize their eggs) with respect to genetic similarity. There was no difference in a female's relatedness to the male in whose territory she resided and her average relatedness to all other males. Neither was there a difference in the relatedness of the male that sired a female's offspring and the female's average relatedness to all other males. There was no evidence of a cost to mating with a more genetically similar mate, because offspring survival was not influenced by degree of inbreeding or outbreeding. However, females that were more outbred had fewer offspring survive. In this small population there are two possible explanations for the decreased fitness associated with outbreeding. First, the species may have an evolutionary history of inbreeding and thus may be susceptible to outbreeding depression. Second. as fitter individuals produce more offspring, these offspring have an increased probability of breeding with relatives, leading to an indirect relationship between fitness and outbreeding.     

Microsatellite diversity and fitness in stranded juvenile harp seals (Phoca groenlandica)

A positive relationship between genetic diversity at neutral markers and juvenile survival has been demonstrated for many vertebrate populations, although the correlation is typically weak and the explanation for it remains controversial. We assessed variation at 9-12 microsatellite loci in 65 juvenile harp seals (Phoca groenlandica) that stranded in poor condition around Long Island, NY, from 2001 to 2004. Compared with seals that died, surviving individuals had slightly higher measures of mean d(2), which reflects the size difference between alleles within an individual and provides an index of outbreeding. In contrast, there were no significant differences between survivors and nonsurvivors in heterozygosity or estimates of internal relatedness. This pattern is attributed to the fact that these microsatellite markers were exceptionally variable in this species (9-22 alleles per locus), and all individuals were heterozygous at most loci. Under these circumstances, mean d(2) may provide a powerful measure for assessing diversity-fitness correlations.     

Multigenerational hybridisation and its consequences for maternal effects in Atlantic salmon

Outbreeding between segregating populations can be important from an evolutionary, conservation and economical-agricultural perspective. Whether and how outbreeding influences maternal effects in wild populations has rarely been studied, despite both the prominent maternal influence on early offspring survival and the known presence of fitness effects resulting from outbreeding in many taxa. We studied several traits during the yolk-feeding stage in multigenerational crosses between a wild and a domesticated Atlantic salmon (Salmo salar) population up to their third-generation hybrid in a common laboratory environment. Using cross-means analysis, we inferred that maternal additive outbreeding effects underlie most offspring traits but that yolk mass also underlies maternal dominant effects. As a consequence of the interplay between additive and dominant maternally controlled traits, offspring from first-generation hybrid mothers expressed an excessive proportion of residual yolk mass, relative to total mass, at the time of first feeding. Their residual yolk mass was 23–97% greater than those of other crosses and 31% more than that predicted by a purely additive model. Offspring additive, epistatic and epistatic offspring-by-maternal outbreeding effects appeared to further modify this largely maternally controlled cross-means pattern, resulting in an increase in offspring size with the percentage of domesticated alleles. Fitness implications remain elusive because of unknown phenotype-by-environment interactions. However, these results suggest how mechanistically co-adapted genetic maternal control on early offspring development can be disrupted by the effects of combining alleles from divergent populations. Complex outbreeding effects at both the maternal and offspring levels make the prediction of hybrid phenotypes difficult.     

Paternal levels of DNA damage in spermatozoa and maternal parity influence offspring mortality in an endangered ungulate

Understanding which factors influence offspring mortality rates is a major challenge since it influences population dynamics and may constrain the chances of recovery among endangered species. Most studies have focused on the effects of maternal and environmental factors, but little is known about paternal factors. Among most polygynous mammals, males only contribute the haploid genome to their offspring, but the possibility that sperm DNA integrity may influence offspring survival has not been explored. We examined several maternal, paternal and individual factors that may influence offspring survival in an endangered species (Gazella cuvieri). Levels of sperm DNA damage had the largest impact upon offspring mortality rates, followed by maternal parity. In addition, there was a significant interaction between these two variables, so that offspring born to primiparous mothers were more likely to die if their father had high levels of sperm DNA damage, but this was not the case among multiparous mothers. Thus, multiparous mothers seem to protect their offspring from the deleterious effects of sperm DNA damage. Since levels of sperm DNA damage seem to be higher among endangered species, more attention should be paid to the impact of this largely ignored factor among the viability of endangered species.     

OPTIMAL OUTCROSSING IN IPOMOPSIS AGGREGAT A: SEED SET AND OFFSPRING FITNEs

Restricted gene flow and localized selection should establish a correlation between physical proximity and genetic similarity in many plant populations. Given this situation, fitness may decline in crosses between nearby plants (inbreeding depression), and in crosses between more widely separated plants (“outbreeding depression”) mostly as a result of disruption of local adaptation. It follows that seed set and offspring fitness may be greatest in crosses over an intermediate “optimal outcrossing distance.” This prediction was supported for Ipomopsis aggregata, a long-lived herbaceous plant pollinated by hummingbirds. In six replicate pollination experiments, mean seed set per flower was higher with an outcrossing distance of 1–10 m than with selfing or outcrossing over 100 m. A similar pattern appeared in the performance of offspring from experimental crosses grown under natural conditions and censused for a seven-year period. Offspring from 10-m crosses had higher survival, greater chance of flowering, and earlier flowering than those from 1-m or 100-m crosses. As a result, 1-m and 100-m offspring achieved only 47% and 68%, respectively, of the lifetime fitness of 10-m offspring. Offspring fitness also declined with planting distance from the seed parent over a range of 1–30 m, so that adaptation to the maternal environment is a plausible mechanism for outbreeding depression. Censuses in a representative I. aggregata population indicated that the herbaceous vegetation changes over a range of 2–150 m, suggesting that there is spatial variation in selection regimes on a scale commensurate with the observed effects of outbreeding depression and planting distance. We discuss the possibility that differences in seed set might in part reflect maternal mate discrimination and emphasize the desirability of measuring offspring fitness under natural conditions in assessing outcrossing effects.     

HETEROZYGOSITY-FITNESS CORRELATIONS REVEALED BY NEUTRAL AND CANDIDATE GENE MARKERS IN ROE DEER FROM A LONG-TERM STUDY

Heterozygosity-fitness correlations (HFCs) are increasingly reported but the underlying mechanisms causing HFCs are generally poorly understood. Here, we test for HFCs in roe deer ( Capreolus capreolus ) using 22 neutral microsatellites widely distributed in the genome and four microsatellites in genes that are potentially under selection. Juvenile survival was used as a proxy for individual fitness in a population that has been intensively studied for 30 years in northeastern France. For 222 juveniles, we computed two measures of genetic diversity: individual heterozygosity ( H ), and mean d ² (relatedness of parental genomes). We found a relationship between genetic diversity and fitness both for the 22 neutral markers and two candidate genes: IGF1 (Insulin-like Growth Factor I) and NRAMP (natural resistance-associated macrophage protein). Statistical evidence and the size of genetic effects on juvenile survival were comparable to those reported for early development and cohort variation, suggesting a substantial influence of genetic components on fitness in this roe deer population. For the 22 neutral microsatellites, a correlation with fitness was revealed for mean d ², but not for H , suggesting a possible outbreeding advantage. This heterosis effect could have been favored by introduction of genetically distant (Hungarian) roe deer to the population in recent times and, possibly, by the structuring of the population into distinct clans. The locus-specific correlations with fitness may be driven by growth rate advantages and resistance to diseases known to exist in the studied population. Our analyses of neutral and candidate gene markers both suggest that the observed HFCs are likely mainly due to linkage with dominant or overdominant loci that affect fitness ("local" effect) rather than to a genome-wide relationship with homozygosity due to inbreeding ("general" effect).     

Host selection by the pine processionary moth enhances larval performance: An experiment

The development of a phytophagous insect depends on the nutritional characteristics of plants on which it feeds. Offspring from different females, however, may vary in their ability to develop in different host species and therefore females should place their eggs on host plants that result in the highest performance for the insect offspring. Causes underlying the predicted relationships between host selection and offspring performance may be: (1) a genetic association between larval ability to exploit particular hosts and the female insect's host preference; and (2) phenotypic plasticity of larvae that may be due to (a) maternal effects (e.g. differential investment in eggs) or (b) diet. In this work, we analyse the performance (i.e. hatching success and larval size and mortality) of the pine processionary (Thaumetopoea pityocampa) caterpillar developing in Aleppo (Pinus halepensis) or maritime (Pinus pinaster) pines. Larvae of this moth species do not move from the individual pine selected by the mother for oviposition. By means of cross-fostering experiments of eggs batches and silk nests of larvae between these two pine species, we explored whether phenotypic plasticity of offspring traits or genetic correlations between mother and offspring traits account for variation in developmental characteristics of caterpillars. Our results showed that females preferentially selected Aleppo pine for oviposition. Moreover, the offspring had the highest probability of survival and reached a larger body size in this pine species independently of whether or not batches were experimentally cross-fostered. Notably, the interaction between identity of donor and receiver pine species of larvae nests explained a significant proportion of variance of larval size and mortality, suggesting a role of diet-induced phenotypic plasticity of the hatchlings. These results suggest that both female selection of the more appropriate pine species and phenotypic plasticity of larva explain the performance of pine processionary caterpillars.     

Phenotypic selection and covariation in the life‐history traits of elephant seals: heavier offspring gain a double selective advantage

Early developmental conditions contribute to individual heterogeneity of both phenotypic traits and fitness components, ultimately affecting population dynamics. Although the demographic consequences of ontogenic growth are best quantified using an integrated measure of fitness, most analyses to date have instead studied individual fitness components in isolation. Here, we estimated phenotypic selection on weaning mass in female southern elephant seals Mirounga leonina by analyzing individual‐based data collected between 1986 and 2016 with capture–recapture and matrix projection models. In support of a hypothesis predicting a gradual decrease of weaning mass effects with time since weaning (the replacement hypothesis), we found that the estimated effects of weaning mass on future survival and recruitment probability was of intermediate duration (rather than transient or permanent). Heavier female offspring had improved odds of survival in early life and a higher probability to recruit at an early age. The positive link between weaning mass and recruitment age is noteworthy, considering that pre‐recruitment mortality already imposed a strong selective filter on the population, leaving only the most ‘robust’ individuals to reproduce. The selection gradient on asymptotic population growth rate, a measure of mean absolute fitness, was weaker than selection on first‐year survival and recruitment probabilities. Weaker selection on mean fitness occurs because weaning mass has little impact on adult survival, the fitness component to which the population growth of long‐lived species is most sensitive. These results highlight the need to interpret individual variation in phenotypic traits in a context that considers the demographic pathways between the trait and an inclusive proxy of individual fitness. Although variation in weaning mass do not translate to permanent survival differences among individuals in adulthood, it explains heterogeneity and positive covariation between survival and breeding in early life, which contribute to between‐individual variation in fitness.     

Patterns of parental relatedness and pup survival in the grey seal (Halichoerus grypus)

Levels of parental relatedness can affect offspring survival and susceptibility to disease. We investigated parental relatedness of live and dead Halichoerus grypus pups between and within island populations and between possible causes of mortality. Nine microsatellites were used to calculate internal relatedness (IR) and standardized mean d2. We find that pups with higher than average levels of IR have significantly lower survival and that this varied between island populations and that certain loci contributed to the effect more than others. Although, there were no significant differences between causes of mortality, peritonitis, infection and stillborn had the highest levels of IR. These results provide evidence that parental relatedness is an important determinant of pre-weaning pup survival in the grey seal and that this may vary with cause of mortality given a larger sample size.     

Evolve II: A computer model of an evolving ecosystem

The Evolve II program is a model of an ecosystem in which organisms are allowed to evolve. Organisms are subject to a changeable environment and competition from other organisms for a limited food supply. The gene structure may change through mutation. A feature of Evolve II is that the magnitude of phenotypic change resulting from mutation is itself a property of the gene. The system was studied under a number of environmental variation schemes. We report three significant findings. Two species (lineages with distinctly different survival strategies) evolved and coexisted in the same environmental conditions. Organisms developed a resistance to phenotypic change in response to mutation in slowly varying environments. However, traits which favor survival of the individual at the expense of reproduction could in some cases undergo phenotypic change in response to mutation despite the fact that this did not favor the survival of the offspring. This demonstrates that gene structures can evolve which are advantageous from the standpoint of the lineage, but not advantageous from the standpoint of individual offspring.     

Variation due to maternal age in the onion root maggot,Hylemya antiqua (Meigen)

A variety of characteristics were studied in populations of the onion root maggot, Hylemya antiqua, to determine the extent to which the age of the female producing a batch of eggs might affect heterogeneity in her offspring. Early-born offspring were found to be reproductively most successful. They had the longest mean expectation of life, a relatively low mortality rate into mid-life, the highest average fecundity, the highest mean rate of egg production, and they produced the highest percentage of female offspring. On the other hand, mid-born offspring were hardiest. They also had a long mean expectation of life, comparable to that for early-born offspring but, in addition, they had the lowest sustained mortality rate and the greatest ability to survive food stress. Their robustness may have been acquired at the expense of certain reproductive capabilities however, in that their average fecundity was lower than their early-born siblings', and their rate of egg production was more variable. Late-born offspring were the most mortality-prone and the least fecund, but they had the shortest mean generation time. These differences must be evaluated in terms of their effects on the rate of natural increase and on population growth. Maternal age was found to influence pupal size, although the provenance groups differed significantly from each other and, in fact, showed opposite trends within their respective cohort groups. Size itself, and the ability of a species to alter the size at which it pupates, may be characteristics which have evolved to fit a specific set of environmental conditions.     

Multigenerational outbreeding effects in Chinook salmon (Oncorhynchus tshawytscha)

Outbreeding, mating between genetically divergent individuals, may result in negative fitness consequences for offspring via outbreeding depression. Outbreeding effects are of notable concern in salmonid research as outbreeding can have major implications for salmon aquaculture and conservation management. We therefore quantified outbreeding effects in two generations (F₁ hybrids and F₂ backcrossed hybrids) of Chinook salmon (Oncorhynchus tshawytscha) derived from captively-reared purebred lines that had been selectively bred for differential performance based on disease resistance and growth rate. Parental lines were crossed in 2009 to create purebred and reciprocal hybrid crosses (n = 53 families), and in 2010 parental and hybrid crosses were crossed to create purebred and backcrossed hybrid crosses (n = 66 families). Although we found significant genetic divergence between the parental lines (F_ST = 0.130), reciprocal F₁ hybrids showed no evidence of outbreeding depression (hybrid breakdown) or favorable heterosis for weight, length, condition or survival. The F₂ backcrossed hybrids showed no outbreeding depression for a suite of fitness related traits measured from egg to sexually mature adult life stages. Our study contributes to the current knowledge of outbreeding effects in salmonids and supports the need for more research to better comprehend the mechanisms driving outbreeding depression.     

Phenotypic plasticity in growth and fecundity induced by strong population fluctuations affects reproductive traits of female fish

Fish are known for their high phenotypic plasticity in life‐history traits in relation to environmental variability, and this is particularly pronounced among salmonids in the Northern Hemisphere. Resource limitation leads to trade‐offs in phenotypic plasticity between life‐history traits related to the reproduction, growth, and survival of individual fish, which have consequences for the age and size distributions of populations, as well as their dynamics and productivity. We studied the effect of plasticity in growth and fecundity of vendace females on their reproductive traits using a series of long‐term incubation experiments. The wild parental fish originated from four separate populations with markedly different densities, and hence naturally induced differences in their growth and fecundity. The energy allocation to somatic tissues and eggs prior to spawning served as a proxy for total resource availability to individual females, and its effects on offspring survival and growth were analyzed. Vendace females allocated a rather constant proportion of available energy to eggs (per body mass) despite different growth patterns depending on the total resources in the different lakes; investment into eggs thus dictated the share remaining for growth. The energy allocation to eggs per mass was higher in young than in old spawners and the egg size and the relative fecundity differed between them: Young females produced more and smaller eggs and larvae than old spawners. In contrast to earlier observations of salmonids, a shortage of maternal food resources did not increase offspring size and survival. Vendace females in sparse populations with ample resources and high growth produced larger eggs and larvae. Vendace accommodate strong population fluctuations by their high plasticity in growth and fecundity, which affect their offspring size and consequently their recruitment and productivity, and account for their persistence and resilience in the face of high fishing mortality.     

Outbreeding depression and breeding system evolution in small,remnant populations of <Emphasis Type="Italic">Primula vulgaris</Emphasis>: consequences for genetic rescue

Many species suffer from anthropogenic habitat fragmentation. The resulting small and isolated populations are more prone to extinction due to, amongst others, genetic erosion, inbreeding depression and Allee-effects. Genetic rescue can help mitigate such problems, but might result in outbreeding depression. We evaluated offspring fitness after selfing and outcrossing within and among three very small and isolated remnant populations of the heterostylous plant Primula vulgaris. We used greenhouse-grown offspring from these populations to test several fitness components. One population was fixed for the pin-morph, and was outcrossed with another population in the field to obtain seeds. Genetic diversity of parent and offspring populations was studied using microsatellites. Morph and population-specific heterosis, inbreeding and outbreeding depression were observed for fruit and seed set, seed weight and cumulative fitness. Highest fitness was observed in the field-outcrossed F1-population, which also showed outbreeding depression following subsequent between-population (back)crossing. Despite outbreeding depression, fitness was still relatively high. Inbreeding coefficients indicated that the offspring were more inbred than their parent populations. Offspring heterozygosity and inbreeding coefficients correlated with observed fitness. One population is evolving homostyly, showing a thrum morph with an elongated style and high autonomous fruit and seed set. This has important implications for conservation strategies such as genetic rescue, as the mating system will be altered by the introduction of homostyles.     

Consequences of outbreeding on phenotypic plasticity in Drosophila mercatorum wings

A multivariate morphometric investigation was conducted on wings of two parthenogenetic Drosophila mercatorum strains and offspring (F1) of crosses between these parthenogenetic strains with highly inbred sexual individuals of the same species. The parental flies and F1 offspring were reared at three different temperatures: 20, 25, or 28°C. This design allows a comparison of completely homozygous individuals (parental generation) with identical heterozygote offspring (F1), which makes an analysis of phenotypic plasticity of morphometric traits possible, without a potentially confounding effect of genotype-environment interactions, which can increase the phenotypic variability. The same pattern of phenotypic plasticity of wing size between the homozygous parental strains and the heterozygous offspring was found in both strains with an apparent heterotic effect for wing size in the F1 at 25°C. At 20 and 28°C flies from the parental generation had the biggest wings. Phenotypic plasticity of shape was found to be strain dependent. A reduced level of developmental instability (DI) was found in the F1 as compared to the parental strain only in strain 1 reared at 20°C for the wing size and 25°C for the wing shape. For all the other treatments higher DI was found in the F1 when the difference was significant, which is suggestive of outbreeding depression. These findings are difficult to interpret since an apparent heterotic effect of size at 25°C is accompanied by higher DI (though not significant in strain 2) and complex changes in wing shape. Hence, we cannot conclude whether outbreeding lowers or increases the capacity to respond to environmental change via plastic responses and via changes of the level of DI. The degree of change of phenotypic plasticity and DI is trait specific, depending on the environment and on the genotypes which are hybridizing. Kristian Krag and Hans Thomsen have contributed equally.     

Genetic rescue persists beyond first-generation outbreeding in small populations of a rare plant

Habitat fragmentation commonly causes genetic problems and reduced fitness when populations become small. Stocking small populations with individuals from other populations may enrich genetic variation and alleviate inbreeding, but such artificial gene flow is not commonly used in conservation owing to potential outbreeding depression. We addressed the role of long-term population size, genetic distance between populations and test environment for the performance of two generations of offspring from between-population crosses of the locally rare plant Ranunculus reptans L. Interpopulation outbreeding positively affected an aggregate measure of fitness, and the fitness superiority of interpopulation hybrids was maintained in the second offspring (F2) generation. Small populations benefited more strongly from interpopulation outbreeding. Genetic distance between crossed populations in neutral markers or quantitative characters was not important. These results were consistent under near-natural competition-free and competitive conditions. We conclude that the benefits of interpopulation outbreeding are likely to outweigh potential drawbacks, especially for populations that suffer from inbreeding.     

Mean d(2) and divergence time: transformations and standardizations

The fitness consequences of inbreeding and outbreeding have intrigued biologists for a long time. Recently a measure of relatedness of parental haplotypes has been proposed called mean d(2). This measure is based on a stepwise mutational process and therefore is tailored to microsatellite genetic markers. Theoretical work suggests that mean d(2) typically is less suited for measuring fitness consequences due to close inbreeding rather than heterozygosity. However, mean d(2) may be more appropriate than heterozygosity for measuring divergence times over longer time scales and thus for detecting outbreeding depression. Here, simulations are used to (1). identify appropriate standardization coefficients and transformations for mean d(2), and (2). evaluate mean d(2) as a measure of divergence time of parental lineages over time scales up to 10000 generations. Results show that mean d(2) is a linear predictor of divergence time. The coefficient of variation of mean d(2) approaches a constant value with increasing divergence time and therefore logarithm transformation is appropriate to restore homoscedasticity. When mutation rates and sizes are known for each locus they can be incorporated into a standardization coefficient to increase the precision of mean d(2). As few as 10 loci can explain more than 70% of the variation in divergence time between lineages. While heterozygosity outperforms mean d(2) at detecting differences in divergence time over relative short time periods (or=1000 generations). However, gene flow of as little as 1% per generation can significantly reduce the ability of either mean d(2) or heterozygosity to estimate divergence time.     

Heterosis and outbreeding depression in descendants of natural immigrants to an inbred population of song sparrows (Melospiza melodia)

We studied heterosis and outbreeding depression among immigrants and their descendants in a population of song sparrows on Mandarte Island, Canada. Using data spanning 19 generations, we compared survival, seasonal reproductive success, and lifetime reproductive success of immigrants, natives (birds with resident-hatched parents and grandparents), and their offspring (F1s, birds with an immigrant and a native parent, and F2s, birds with an immigrant grandparent and resident-hatched grandparent in each of their maternal and paternal lines). Lifetime reproductive success of immigrants was no worse than that of natives, but other measures of performance differed in several ways. Immigrant females laid later and showed a tendency to lay fewer clutches, but had relatively high success raising offspring per egg produced. The few immigrant males survived well but were less likely to breed than native males of the same age that were alive in the same year. Female F1s laid earlier than expected based on the average for immigrant and native females, and adult male F1s were more likely to breed than expected based on the average for immigrant and native males. The performance differences between immigrant and native females and between F1s and the average of immigrants and natives are consistent with the hypothesis that immigrants were disadvantaged by a lack of site experience and that immigrant offspring benefited from heterosis. However, we could not exclude the possibility that immigrants had a different strategy for optimizing reproductive success or that they experienced ecological compensation for life-history parameters. For example, the offspring of immigrants may have survived well because immigrants laid later and produced fewer clutches, thereby raising offspring during a period of milder climatic conditions. Although sample sizes were small, we found large performance differences between F1s and F2s, which suggested that either heterosis was associated with epistasis in F1s, that F2s experienced outbreeding depression, or that both phenomena occurred. These findings indicate that the performance of dispersers may be affected more by fine-scale genetic differentiation than previously assumed in this and comparable systems.     

Lower fitness of hatchery and hybrid rainbow trout compared to naturalized populations in Lake Superior tributaries

We have documented an early life survival advantage by naturalized populations of anadromous rainbow trout Oncorhynchus mykiss over a more recently introduced hatchery population and outbreeding depression resulting from interbreeding between the two strains. We tested the hypothesis that offspring of naturalized and hatchery trout, and reciprocal hybrid crosses, survive equally from fry to age 1+ in isolated reaches of Lake Superior tributary streams in Minnesota. Over the first summer, offspring of naturalized females had significantly greater survival than offspring of hatchery females in three of four comparisons (two streams and 2 years of stocking). Having an entire naturalized genome, not just a naturalized mother, was important for survival over the first winter. Naturalized offspring outperformed all others in survival to age 1+ and hybrids had reduced, but intermediate, survival relative to the two pure crosses. Averaging over years and streams, survival relative to naturalized offspring was 0.59 for hybrids with naturalized females, 0.37 for the reciprocal hybrids, and 0.21 for hatchery offspring. Our results indicate that naturalized rainbow trout are better adapted to the conditions of Minnesota's tributaries to Lake Superior so that they outperform the hatchery-propagated strain in the same manner that many native populations of salmonids outperform hatchery or transplanted fish. Continued stocking of the hatchery fish may conflict with a management goal of sustaining the naturalized populations.