Climate change vulnerability of Arctic char across Scandinavia.

Climate change is anticipated to cause species to shift their ranges upward and poleward, yet space for tracking suitable habitat conditions may be limited for range-restricted species at the highest elevations and latitudes of the globe. Consequently, range-restricted species inhabiting Arctic freshwater ecosystems, where global warming is most pronounced, face the challenge of coping with changing abiotic and biotic conditions or risk extinction. Here, we use an extensive fish community and environmental dataset for 1762 lakes sampled across Scandinavia (mid-1990s) to evaluate the climate vulnerability of Arctic char (Salvelinus alpinus), the world's most cold-adapted and northernly distributed freshwater fish. Machine learning models show that abiotic and biotic factors strongly predict the occurrence of Arctic char across the region with an overall accuracy of 89 percent. Arctic char is less likely to occur in lakes with warm summer temperatures, high dissolved organic carbon levels (i.e., browning), and presence of northern pike (Esox lucius). Importantly, climate warming impacts are moderated by habitat (i.e., lake area) and amplified by the presence of competitors and/or predators (i.e., northern pike). Climate warming projections under the RCP8.5 emission scenario indicate that 81% of extant populations are at high risk of extirpation by 2080. Highly vulnerable populations occur across their range, particularly near the southern range limit and at lower elevations, with potential refugia found in some mountainous and coastal regions. Our findings highlight that range shifts may give way to range contractions for this cold-water specialist, indicating the need for pro-active conservation and mitigation efforts to avoid the loss of Arctic freshwater biodiversity.

Brown trout (Salmo trutta L. 1758) and Arctic charr [Salvelinus alpinus (L. 1758)] display different marine behaviour and feeding strategies in sympatry.

Brown trout (Salmo trutta L. 1758) and Arctic charr [Salvelinus alpinus (L. 1758)] tagged with acoustic transmitters migrated from fresh water to the sea mainly in May and June, but with large individual variation in migration timing. For S. trutta, large individuals (42-86 cm total length) migrated earlier in the season than small individuals (18-27 cm). For S. alpinus, no such pattern was found, likely because of the small size range of tagged fish (28-41 cm). S. trutta stayed longer at sea than S. alpinus (average 2 vs. 1 month). Early migrants of S. trutta stayed for a shorter period at sea than late migrants, whereas no such pattern was observed for S. alpinus. Large S. trutta moved quickly away from the river and spent average 3 days to reach a receiver line 20 km from the river mouth, whereas small S. trutta and S. alpinus migrating that far spent 2-3 weeks on the same distance. S. trutta utilized the entire fjord system and had a greater proportion of long-distance migrants (>20 km, 78% and 59% of large and small S. trutta, respectively) than S. alpinus (29%). S. alpinus mostly stayed in the inner fjord areas, and none were recorded in the outermost part of the fjord. The difference in the use of marine areas may be caused by variation in prey choice and spatial distribution of the preferred prey groups. Stable isotope analysis showed that S. trutta had been feeding at a higher trophic level than S. alpinus. S. trutta had mainly fed on marine fish and shrimps, whereas S. alpinus had large proportions of freshwater invertebrates in the diet, suggesting that the estuary with benthos and amphipods drifting from the river was an important feeding habitat for S. alpinus. In conclusion, major differences in habitat use, migration patterns and feeding strategies were found between sympatric anadromous S. trutta and S. alpinus while at sea.

Salinity-induced transcriptome profiles in marine and freshwater threespine stickleback after an abrupt 6-hour exposure.

Saltwater and freshwater environments have opposing physiological challenges, yet, there are fish species that are able to enter both habitats during short time spans, and as individuals they must therefore adjust quickly to osmoregulatory contrasts. In this study, we conducted an experiment to test for plastic responses to abrupt salinity changes in two populations of threespine stickleback, Gasterosteus aculeatus, representing two ecotypes (freshwater and ancestral saltwater). We exposed both ecotypes to abrupt native (control treatment) and non-native salinities (0‰ and 30‰) and sampled gill tissue for transcriptomic analyses after 6 h of exposure. To investigate genomic responses to salinity, we analyzed four different comparisons; one for each ecotype (in their control and exposure salinity; (1) and (2), one between ecotypes in their control salinity (3), and the fourth comparison included all transcripts identified in (3) that did not show any expressional changes within ecotype in either the control or the exposed salinity (4)). Abrupt salinity transfer affected the expression of 10 and 1530 transcripts for the saltwater and freshwater ecotype, respectively, and 1314 were differentially expressed between the controls, including 502 that were not affected by salinity within ecotype (fixed expression). In total, these results indicate that factors other than genomic expressional plasticity are important for osmoregulation in stickleback, due to the need for opposite physiological pathways to survive the abrupt change in salinity.

Ecological ramifications of adaptation to size-selective mortality.

Size-selective mortality due to harvesting is a threat to numerous exploited species, but how it affects the ecosystem remains largely unexplored. Here, we used a pond mesocosm experiment to assess how evolutionary responses to opposite size-selective mortality interacted with the environment (fish density and light intensity used as a proxy of resource availability) to modulate fish populations, prey community composition and ecosystem functions. We used medaka (Oryzias latipes) previously selected over 10 generations for small size (harvest-like selection; small-breeder line) or large size (large-breeder line), which displayed slow somatic growth and early maturity or fast somatic growth and late maturity, respectively. Large-breeder medaka produced more juveniles, which seemed to grow faster than small-breeder ones but only under high fish density. Additionally, large-breeder medaka had an increased impact on some benthic prey, suggesting expanded diet breadth and/or enhanced foraging abilities. As a consequence, increased light stimulated benthic algae biomass only in presence of large-breeder medaka, which were presumably better at controlling benthic grazers. Aggregated effect sizes at the community and ecosystem levels revealed that the ecological effects of medaka evolution were of similar magnitude to those induced by the environment and fish introduction. These findings indicate the important environmental dependency of evolutionary response to opposite size-selective mortality on higher levels of biological organizations.

Movement patterns of temperate wrasses (Labridae) within a small marine protected area.

The movement patterns of three commercially important wrasse (Labridae) species inside a small marine protected area (~ 0.15 km2 ) on the west coast of Norway were analysed over a period of 21 months. The mean distance between capture and recapture locations varied between 10 and 187 m, and was species and season specific. The extent of movement was not related to body size or sex. These results imply that a network of small strategically located marine protected areas can be used as management tools to protect wrasses from size- and sex-selective fishing mortality.

Marine protected areas rescue a sexually selected trait in European lobster.

Marine protected areas (MPAs) are increasingly implemented worldwide to maintain and restore depleted populations. However, despite our knowledge on the myriad of positive responses to protection, there are few empirical studies on the ability to conserve species' mating patterns and secondary sexual traits. In male European lobsters (Homarus gammarus), the size of claws relative to body size correlates positively with male mating success and is presumably under sexual selection. At the same time, an intensive trap fishery exerts selection against large claws in males. MPAs could therefore be expected to resolve these conflicting selective pressures and preserve males with large claws. We explored this hypothesis by contrasting claw size of males and females in three pairs of MPAs and nearby fished areas in southern Norway. By finding that male lobsters have up to 8% larger claws inside MPAs compared to similarly sized males in fished areas, our study provides evidence that MPAs rescue a secondary sexual trait. Recovery from harvest selection acting on claws is the most likely explanation; however, the higher abundance of lobster inside MPAs does not rule out a plastic response on claw size due to increased competition. Regardless of the underlying cause, our study demonstrates (a) the value of protected areas as a management tool for mitigating fisheries-induced evolution and (b) that MPAs help maintaining the scope for sexual selection in populations with vulnerable life histories and complex mating system.

Age of European silver eels during a period of declining abundance in Norway.

The European eel (Anguilla anguilla) is critically endangered throughout its range. Knowledge about age distribution of future spawners (silver eels) is essential to monitor the status and contribute to the recovery of this species. Determination of age in anguillid eels is challenging, especially in eels from the northern part of the distribution area where growth is slow and age at maturation can be up to 30 years or more. Eels from the river Imsa in Norway have been monitored since 1975, and this reference time series has been used to assess the stock at the European level. Population dynamics in this catchment were analyzed during the late 1980s by estimating ages on whole cleared otoliths. However, techniques for revealing annual increments on otoliths have evolved over the years sometimes yielding significant differences in age estimates. In this study, the historical otolith data were reanalyzed using a grinding and polishing method rather than reading the whole otolith. The new age estimates were considerably higher than the previous ones, sometimes by up to 29 years. Since the 1980s, mean age of silver eels only slightly increased (from 19 to 21 years in the 2010s). This was mainly due to the disappearance of younger silver eels (<15 years) in the 2010s. The new age estimates agreed with the steep decline in recruitment which occurred in the late 1980s in the Imsa catchment. Mean growth (30 mm/year, min-max: 16-64 mm/year) has not changed since the 1980s, although density in the catchment has decreased. Revealing and reading age of slow-growing eels remain a challenge but adding a measure of otolith reading uncertainty may improve age data collection and contribute to recovery measures for this species.

Size- and stage-dependence in cause-specific mortality of migratory brown trout.

Evidence-based management of natural populations under strong human influence frequently requires not only estimates of survival but also knowledge about how much mortality is due to anthropogenic vs. natural causes. This is the case particularly when individuals vary in their vulnerability to different causes of mortality due to traits, life history stages, or locations. Here, we estimated harvest and background (other cause) mortality of landlocked migratory salmonids over half a century. In doing so, we quantified among-individual variation in vulnerability to cause-specific mortality resulting from differences in body size and spawning location relative to a hydropower dam. We constructed a multistate mark-recapture model to estimate harvest and background mortality hazard rates as functions of a discrete state (spawning location) and an individual time-varying covariate (body size). We further accounted for among-year variation in mortality and migratory behaviour and fit the model to a unique 50-year time series of mark-recapture-recovery data on brown trout (Salmo trutta) in Norway. Harvest mortality was highest for intermediate-sized trout, and outweighed background mortality for most of the observed size range. Background mortality decreased with body size for trout spawning above the dam and increased for those spawning below. All vital rates varied substantially over time, but a trend was evident only in estimates of fishers' reporting rate, which decreased from over 50% to less than 10% throughout the study period. We highlight the importance of body size for cause-specific mortality and demonstrate how this can be estimated using a novel hazard rate parameterization for mark-recapture models. Our approach allows estimating effects of individual traits and environment on cause-specific mortality without confounding, and provides an intuitive way to estimate temporal patterns within and correlation among different mortality sources.

Stoichiometric consequences of size-selective mortality: An experimental test using the Japanese medaka (Oryzias latipes).

The determinants of intraspecific stoichiometric variation remain difficult to elucidate due to their multiple origins (e.g. genetic vs. environmental) and potential interactive effects. We evaluated whether two size-selected lines of medaka (Oryzias latipes) with contrasted life-history strategies (small- and large-breeder lines with slow growth and early maturity vs. fast growth and late maturity) differed in their organismal stoichiometry (percentage and ratios of carbon [C], nitrogen [N] and phosphorus [P]) in a mesocosm experiment. We also tested how size-selection interacted with environmental conditions (i.e. two levels of fish density and light intensity), body condition and sex. Results showed that large-breeder fish were significantly N-enriched compared to small-breeders, while the two size-selected lines did not differ in body P composition. Size-selection interacted with density - high density only affected small-breeders leading to decreasing %C and C: N - and with sex - large-breeder females had higher %C and C:N values than large-breeder males. Finally, C:P and N:P ratios increased with body condition due to decreasing %P. Overall, our results show that the ecological consequences of size-selective mortality extend to organismal stoichiometry and may, from there, change nutrient cycling and ecosystem functioning.

Effects of size- and sex-selective harvesting: An integral projection model approach.

Harvesting is often size-selective, and in species with sexual size dimorphism, it may also be sex-selective. A powerful approach to investigate potential consequences of size- and/or sex-selective harvesting is to simulate it in a demographic population model. We developed a population-based integral projection model for a size- and sex-structured species, the commonly exploited pike (Esox lucius). The model allows reproductive success to be proportional to body size and potentially limited by both sexes. We ran all harvest simulations with both lower size limits and slot limits, and to quantify the effects of selective harvesting, we calculated sex ratios and the long-term population growth rate (λ). In addition, we quantified to what degree purely size-selective harvesting was sex-selective, and determined when λ shifted from being female to male limited under size- and sex-selective harvesting. We found that purely size-selective harvest can be sex-selective, and that it depends on the harvest limits and the size distributions of the sexes. For the size- and sex-selective harvest simulations, λ increased with harvest intensity up to a threshold as females limited reproduction. Beyond this threshold, males became the limiting sex, and λ decreased as more males were harvested. The peak in λ, and the corresponding sex ratio in harvest, varied with both the selectivity and the intensity of the harvest simulation. Our model represents a useful extension of size-structured population models as it includes both sexes, relaxes the assumption of female dominance, and accounts for size-dependent fecundity. The consequences of selective harvesting presented here are especially relevant for size- and sex-structured exploited species, such as commercial fisheries. Thus, our model provides a useful contribution toward the development of more sustainable harvesting regimes.

Harvesting changes mating behaviour in European lobster.

Removing individuals from a wild population can affect the availability of prospective mates and the outcome of competitive interactions, with subsequent effects on mating patterns and sexual selection. Consequently, the rate of harvest-induced evolution is predicted to be strongly dependent on the strength and dynamics of sexual selection, yet there is limited empirical knowledge on the interplay between selective harvesting and the mating systems of exploited species. In this study, we used genetic parentage assignment to compare mating patterns of the highly valued and overexploited European lobster (Homarus gammarus) in a designated lobster reserve and nearby fished area in southern Norway. In the area open to fishing, the fishery is regulated by a closed season, a minimum legal size and a ban on the harvest of egg-bearing females. Due to the differences in size and sex-specific fishing mortality between the two areas, males and females are of approximately equal average size in the fished area, whereas males tend to be larger in the reserve. Our results show that females would mate with males larger than their own body size, but the relative size difference was significantly larger in the reserve. Sexual selection acted positively on both body size and claw size in males in the reserve, while it was nonsignificant in fished areas. This strongly suggests that size truncation of males by fishing reduces the variability of traits that sexual selection acts upon. If fisheries continue to target large individuals (particularly males) with higher relative reproductive success, the weakening of sexual selection will likely accelerate fisheries-induced evolution towards smaller body size.

Ongoing niche differentiation under high gene flow in a polymorphic brackish water threespine stickleback (Gasterosteus aculeatus) population.

BACKGROUND: Marine threespine sticklebacks colonized and adapted to brackish and freshwater environments since the last Pleistocene glacial. Throughout the Holarctic, three lateral plate morphs are observed; the low, partial and completely plated morph. We test if the three plate morphs in the brackish water Lake Engervann, Norway, differ in body size, trophic morphology (gill raker number and length), niche (stable isotopes; δ15N, δ13C, and parasites (Theristina gasterostei, Trematoda spp.)), genetic structure (microsatellites) and the lateral-plate encoding Stn382 (Ectodysplasin) gene. We examine differences temporally (autumn 2006/spring 2007) and spatially (upper/lower sections of the lake - reflecting low versus high salinity). RESULTS: All morphs belonged to one gene pool. The complete morph was larger than the low plated, with the partial morph intermediate. The number of lateral plates ranged 8-71, with means of 64.2 for complete, 40.3 for partial, and 14.9 for low plated morph. Stickleback δ15N was higher in the lower lake section, while δ13C was higher in the upper section. Stickleback isotopic values were greater in autumn. The low plated morph had larger variances in δ15N and δ13C than the other morphs. Sticklebacks in the upper section had more T. gasterostei than in the lower section which had more Trematoda spp. Sticklebacks had less T. gasterostei, but more Trematoda spp. in autumn than spring. Sticklebacks with few and short rakers had more T. gasterostei, while sticklebacks with longer rakers had more Trematoda. spp. Stickleback with higher δ15N values had more T. gasterostei, while sticklebacks with higher δ15N and δ13C values had more Trematoda spp. The low plated morph had fewer Trematoda spp. than other morphs. CONCLUSIONS: Trait-ecology associations may imply that the three lateral plate morphs in the brackish water lagoon of Lake Engervann are experiencing ongoing divergent selection for niche and migratory life history strategies under high gene flow. As such, the brackish water zone may generally act as a generator of genomic diversity to be selected upon in the different environments where threespine sticklebacks can live.

Individual heterogeneity and early life conditions shape growth in a freshwater top predator.

Body size can have profound impacts on survival, movement, and reproductive schedules shaping individual fitness, making growth a central process in ecological and evolutionary dynamics. Realized growth is the result of a complex interplay between life history schedules, individual variation, and environmental influences. Integrating all of these aspects into growth models is methodologically difficult, depends on the availability of repeated measurements of identifiable individuals, and consequently represents a major challenge in particular for natural populations. Using a unique 30-yr time series of individual length measurements inferred from scale year rings of wild brown trout, we develop a Bayesian hierarchical model to estimate individual growth trajectories in temporally and spatially varying environments. We reveal a gradual decrease in average juvenile growth, which has carried over to adult life and contributed to decreasing sizes observed at the population level. Commonly studied environmental drivers like temperature and water flow did not explain much of this trend and overall persistent and among-year individual variation dwarfed temporal variation in growth patterns. Our model and results are relevant to a wide range of questions in ecology and evolution requiring a detailed understanding of growth patterns, including conservation and management of many size-structured populations.

Timing and pattern of annual silver eel migration in two European watersheds are determined by similar cues.

Many animals perform long-distance migrations in order to maximize lifetime reproductive success. The European eel migrates several thousand kilometers between their feeding habitats in continental waters (fresh-, brackish, and sea water) and their spawning area in the Sargasso Sea. Eels residing in freshwaters usually initiate their spawning migration as silver eels during autumn, triggered by diverse environmental cues. We analyzed the time series of silver eel downstream migration in Burrishoole, Ireland (1971-2015), and Imsa, Norway (1975-2015), to examine factors regulating the silver eel migration from freshwater to the sea. The migration season (90% of the run) generally lasted from 1 August to 30 November. Environmental factors acting in the months before migration impacted timing and duration of migration, likely through influencing the internal processes preparing the fish for migration. Once the migration had started, environmental factors impacted the day-to-day variation in number of migrants, apparently stimulating migration among those eels ready for migration. Both the day-to-day variation in the number of migrants and the onset of migration were described by nearly identical models in the two rivers. Variables explaining day-to-day variation were all associated with conditions that may minimize predation risk; number of migrants was reduced under a strong moon and short nights and increased during high and increasing water levels. Presence of other migrants stimulated migration, which further indicates that silver eel migration has evolved to minimize predation risk. The onset of migration was explained mainly by water levels in August. The models for duration of the migration season were less similar between the sites. Thus, the overall migration season seems governed by the need to reach the spawning areas in a synchronized manner, while during the actual seaward migration, antipredator behavior seems of overriding importance.

The temporal window of ecological adaptation in postglacial lakes: a comparison of head morphology, trophic position and habitat use in Norwegian threespine stickleback populations.

BACKGROUND: Studying how trophic traits and niche use are related in natural populations is important in order to understand adaptation and specialization. Here, we describe trophic trait diversity in twenty-five Norwegian freshwater threespine stickleback populations and their putative marine ancestor, and relate trait differences to postglacial lake age. By studying lakes of different ages, depths and distance to the sea we examine key environmental variables that may predict adaptation in trophic position and habitat use. We measured trophic traits including geometric landmarks that integrated variation in head shape as well as gillraker length and number. Trophic position (Tpos) and niche use (α) were estimated from stable isotopes (δ(13)C, δ(15)N). A comparison of head shape was also made with two North American benthic-limnetic species pairs. RESULTS: We found that head shape differed between marine and freshwater sticklebacks, with marine sticklebacks having more upturned mouths, smaller eyes, larger opercula and deeper heads. Size-adjusted gillraker lengths were larger in marine than in freshwater stickleback. Norwegian sticklebacks were compared on the same head shape axis as the one differentiating the benthic-limnetic North American threespine stickleback species pairs. Here, Norwegian freshwater sticklebacks with a more "limnetic head shape" had more and longer gillrakers than sticklebacks with "benthic head shape". The "limnetic morph" was positively associated with deeper lakes. Populations differed in α (mean ± sd: 0.76 ± 0.29) and Tpos (3.47 ± 0.27), where α increased with gillraker length. Larger fish had a higher Tpos than smaller fish. Compared to the ecologically divergent stickleback species pairs and solitary lake populations in North America, Norwegian freshwater sticklebacks had similar range in Tpos and α values, but much less trait divergences. CONCLUSIONS: Our results showed trait divergences between threespine stickleback in marine and freshwater environments. Freshwater populations diverged in trophic ecology and trophic traits, but trophic ecology was not related to the elapsed time in freshwater. Norwegian sticklebacks used the same niches as the ecologically divergent North American stickleback species pairs. However, as trophic trait divergences were smaller, and not strongly associated with the ecological niche, ecological adaptations along the benthic-limnetic axis were less developed in Norwegian sticklebacks.

Lateral plate number in low-plated threespine stickleback: a study of plasticity and heritability.

In the threespine stickleback Gasterosteus aculeatus model system, phenotypes are often classified into three morphs according to lateral plate number. Morph identity has been shown to be largely genetically determined, but substantial within-morph variation in plate number exists. In this study, we test whether plate number has a plastic component in response to salinity in the low-plated morph using a split-clutch experiment where families were split in two, one half raised in water at 0 and the other at 30 ppt salt. We find a small salinity-induced plastic effect on plate number in an unexpected direction, opposite to what we predicted: Fish raised in freshwater on average have slightly more plates than fish raised in saltwater. Our results confirm that heritability of plate number is high. Additionally, we find that variance in plate number at the family level can be predicted from other family level traits, which might indicate that epistatic interactions play a role in creating the observed pattern of lateral plate number variation.

Genomic signatures of the plateless phenotype in the threespine stickleback.

Understanding the genetic basis of traits involved in adaptive divergence and speciation is one of the most fundamental objectives in evolutionary biology. Toward that end, we look for signatures of extreme plate loss in the genome of freshwater threespine sticklebacks (Gasterosteus aculeatus). Plateless stickleback have been found in only a few lakes and streams across the world; they represent the far extreme of a phenotypic continuum (plate number) that has been studied for years, although plateless individuals have not yet been the subject of much investigation. We use a dense single nucleotide polymorphism dataset made using RADseq to study fish from three freshwater populations containing plateless and low plated individuals, as well as fish from full plated marine populations. Analyses were performed using FastStructure, sliding windows F ST, Bayescan and latent factor mixed models to search for genomic differences between the low plated and plateless phenotypes both within and among the three lakes. At least 18 genomic regions which may contribute to within-morph plate number variation were detected in our low plated stickleback populations. We see no evidence of a selective sweep between low and plateless fish; rather reduction of plate number within the low plated morph seems to be polygenic.

Gyrodactylus spp. diversity in native and introduced minnow (Phoxinus phoxinus) populations: no support for "the enemy release" hypothesis.

BACKGROUND: Translocation of native species and introduction of non-native species are potentially harmful to the existing biota by introducing e.g. diseases, parasites and organisms that may negatively affect the native species. The enemy release hypothesis states that parasite species will be lost from host populations when the host is introduced into new environments. METHODS: We tested the enemy release hypothesis by comparing 14 native and 29 introduced minnow (Phoxinus phoxinus) populations in Norway with regard to the ectoparasitic Gyrodactylus species community and load (on caudal fin). Here, we used a nominal logistic regression on presence/absence of Gyrodactylus spp. and a generalized linear model on the summed number of Gyrodactylus spp. on infected populations, with individual minnow heterozygosity (based on 11 microsatellites) as a covariate. In addition, a sample-based rarefaction analysis was used to test if the Gyrodactylus-species specific load differed between native and introduced minnow populations. An analysis of molecular variance was performed to test for hierarchical population structure between the two groups and to test for signals of population bottlenecks the two-phase model in the Wilcoxon signed-rank test was used. To test for demographic population expansion events in the introduced minnow population, we used the kg-test under a stepwise mutation model. RESULTS: The native and introduced minnow populations had similar species compositions of Gyrodactylus, lending no support to the enemy release hypothesis. The two minnow groups did not differ in the likelihood of being infected with Gyrodactylus spp. Considering only infected minnow populations it was evident that native populations had a significantly higher mean abundance of Gyrodactylus spp. than introduced populations. The results showed that homozygotic minnows had a higher Gyrodactylus spp. infection than more heterozygotic hosts. Using only infected individuals, the two minnow groups did not differ in their mean number of Gyrodactylus spp. However, a similar negative association between heterozygosity and abundance was observed in the native and introduced group. There was no evidence for demographic bottlenecks in the minnow populations, implying that introduced populations retained a high degree of genetic variation, indicating that the number of introduced minnows may have been large or that introductions have been happening repeatedly. This could partly explain the similar species composition of Gyrodactylus in the native and introduced minnow populations. CONCLUSIONS: In this study it was observed that native and introduced minnow populations did not differ in their species community of Gyrodactylus spp., lending no support to the enemy release hypothesis. A negative association between individual minnow host heterozygosity and the number of Gyrodactylus spp. was detected. Our results suggest that the enemy release hypothesis does not necessarily limit fish parasite dispersal, further emphasizing the importance of invasive fish species dispersal control.

Fitness consequences of early life conditions and maternal size effects in a freshwater top predator.

Conditions experienced in early life stages can be an important determinant of individual life histories. In fish, environmental conditions are known to affect early survival and growth, but recent studies have also emphasized maternal effects mediated by size or age. However, the relative sensitivity of the mean fitness (population growth rate λ) to different early life impacts remains largely unexplored. Using a female-based integral projection model (IPM) parameterized from unique long-term demographic data for pike (Esox lucius), we evaluated the relative fitness consequences of different early life impacts, including (i) maternal effects of length on egg weight, potentially affecting offspring (first year) survival, and (ii) effects of temperature on offspring growth and survival. Of the seven vital rates defining the model, offspring survival could not be directly estimated and four scenarios were defined for this rate. Elasticity analyses of the IPM were performed to calculate (i) the total contribution from different lengths to the elasticity of λ to the projection kernel, and (ii) the elasticity of λ to underlying variables of female current length, female offspring length at age 1, and temperature. These elasticities were decomposed into contributions from different vital rates across length. Egg weight increased with female length, as expected, but the effect leveled off for the largest females. However, λ was largely insensitive to this effect, even when egg weight was assumed to have a strong effect on offspring survival. In contrast, λ was sensitive to early temperature conditions through growth and survival. Among mature females, the total elasticity of λ to the projection kernel generally increased with length. The results were robust to a wide range of assumptions. These results suggest that environmental conditions experienced in early life represent a more important driver of mean population growth and fitness of pike than maternal effects of size on offspring survival. We discuss two general mechanisms underlying the weak influence of this maternal effect, suggesting that these may be general for long-lived and highly fecund fishes. This model and results are relevant for the management of long-lived top predators, including many commercially important fish species.

Plastic and Evolutionary Gene Expression Responses Are Correlated in European Grayling (Thymallus thymallus) Subpopulations Adapted to Different Thermal Environments.

Understanding how populations adapt to changing environmental conditions is a long-standing theme in evolutionary biology. Gene expression changes have been recognized as an important driver of local adaptation, but relatively little is known regarding the direction of change and in particular, about the interplay between plastic and evolutionary gene expression. We have previously shown that the gene expression profiles of European grayling (Thymallus thymallus) populations inhabiting different thermal environments include both plastic and evolutionary components. However, whether the plastic and evolutionary responses were in the same direction was not investigated in detail, nor was the identity of the specific genes involved. In this study, we show that the plastic changes in protein expression in response to different temperatures are highly correlated with the evolutionary response in grayling subpopulations adapted to different thermal environments. This finding provides preliminary evidence that the plastic response most likely facilitates adaptation during the early phases of colonization of thermal environments. The proteins that showed significant changes in expression level between warm and cold temperature treatments were mostly related to muscle development, which is consistent with earlier findings demonstrating muscle mass differentiation between cold and warm grayling populations.