Regional differentiation of North American Atlantic salmon at allozyme loci

Allozyme variation was characterised by starch gel electrophoresis at 23 enzyme coding loci and one regulatory locus in Atlantic salmon from 53 rivers in Eastern Canada, encompassing the majority of the species' North American range. Variation among rivers was highly heterogeneous and eight of the 15 polymorphisms showed regionally restricted distributions. Nearest neighbour joining (NJ) analysis and multi‐dimensional scaling suggest six distinct regional groups; Labrador/Ungava, Gulf of Saint Lawrence, Newfoundland (excluding Gulf rivers), the Atlantic shore/Southern Uplands of Nova Scotia, the inner Bay of Fundy, and the outer Bay of Fundy. Approximately 25% of observed genetic variation was distributed among these regions with a weak though significant overall correlation of genetic and geographic distance (Mantel Test, r  = 0·255, P  = 0·005). Collectively, the rivers showed consistent divergence from European populations with strong bootstrap support for the two clusters across loci in the NJ analysis. Mean heterozygosity was 0·061 for both continental groups, but the European population showed more than twice the variation among populations. F _ST values were 0·076 and 0·176 for North America and Europe, respectively, with an overall F _ST of 0·330.     

Population structure of lake‐type and river‐type sockeye salmon in transboundary rivers of northern British Columbia

The population structure of 'lake‐type' and 'river‐type' sockeye salmon Oncorhynchus nerka , primarily in transboundary rivers in northern British Columbia, was examined with a survey of microsatellite variation. Variation at 14 microsatellite loci was surveyed from c . 3000 lake‐type and 3200 river‐type sockeye salmon from 47 populations in six river drainages in British Columbia. The mean F _ST for the 14 microsatellite loci and 47 populations was 0·068, and 0·034 over all river‐type populations. River‐type sockeye salmon were more genetically diverse than lake‐type sockeye salmon, with expected heterozygosity of river‐type sockeye salmon 0·72 and with an average 12·7 alleles observed per locus, whereas expected heterozygosity of lake‐type sockeye salmon was 0·65 with and average 10·5 alleles observed per locus. River drainage of origin was a significant unit of population structure. There was clear evidence of genetic differentiation among river‐type populations of sockeye salmon from different drainages over a broad geographic range in British Columbia.     

Temporal stability of sea louse Lepeophtheirus salmonis Krøyer populations on Atlantic salmon Salmo salar L. of wild, farm and hybrid parentage

Atlantic salmon salmo salar smolts of wild, hybrid and farmed parentage were individually tagged then reared in a sea cage for 8 months. The fish were sampled three times during this period. On all occasions, farmed Atlantic salmon displayed the highest abundance and density of sea lice Lepeophtheirus salmonis , whilst no significant differences were observed between hybrid and wild Atlantic salmon. Percentage variation between the lowest and highest infected groups was as high as 175 and 144% for L. salmonis abundance and density respectively (sample 2). The temporal stability of interindividual sea lice infection levels was investigated pair‐wise between samples using correlation (sample 1 v . 2, 1 v . 3 and 2 v . 3). When calculated using sea louse abundance, correlations ranged from r ² = 0·11, P  < 0·01 to r ² = 0·39, P  < 0·001, but, when the effects of fish size were controlled for by converting abundance to density, all correlations were <  r ² = 0·1. Therefore, these data indicate that a fish's relative infection level in one sample was a weak predictor of its relative infection level in another sample. This suggests that identification of individual Atlantic salmon that display reduced susceptibility to sea lice, may be problematic.     

Inter‐individual differences in rates of routine energy loss and growth in young‐of‐the‐year juvenile Atlantic cod

Inter‐individual differences in rates of routine (non‐feeding) metabolism and growth were evaluated in young‐of‐the‐year (YOY) juvenile Atlantic cod Gadus morhua . Rates of O₂ consumption, CO₂ production and ammonia (TAN) excretion were measured in 64, 25–43 mm standard length ( L _S) YOY growing at different rates (0·27–0·47 mm day⁻¹) in a common rearing tank. Parameter rates ( y ) increased allometrically ( y = a·M^b ) with increasing body mass ( M ) with b ‐values for O₂ production, CO₂ consumption and TAN excretion equal to 0·81, 0·89 and 0·56, respectively. In some cases, residuals from these regressions were significantly negatively correlated to fish growth rate. In no cases did residuals of parameter rates increase with increasing growth rate. These data suggest that, during unfed periods, relatively fast‐growing fish were more metabolically efficient than slower‐growing fish from the same cohort. The fish condition factor, derived from     , also significantly decreased with increasing growth rate. Results indicated differences in both the rates of routine energy loss and the patterns of growth allocation among YOY Atlantic cod. Since these physiological attributes were positively correlated with growth rate, they may be indicative of 'survivors' in field populations.     

Temporal and spatial variation in growth of juvenile Atlantic salmon

Spatial and temporal variation in length‐at‐age and environmental factors affecting variation in growth rate of juvenile Atlantic salmon Salmo salar were studied using data from a long‐term study in the River Stjørdalselva, central Norway. Mean annual instantaneous growth rate among 1+ and 2+year juvenile Atlantic salmon varied between 0·59 and 1·50 g g⁻¹ year⁻¹ and mean instantaneous daily growth rate of young‐of‐the‐year (YOY) varied between 0·013 and 0·033 g g⁻¹ day⁻¹. Between year variation in growth was larger than the within year intra‐watercourse spatial variation. For YOY and 1+year Atlantic salmon, a major part of the observed between year variation in growth rates was explained by variation in mean daily water discharge and spring temperature. For 2+year juvenile Atlantic salmon, mean daily water discharge and cohort density were the only variables to significantly explain variation in growth rates. A large part of the within water‐course spatial variation could not be explained by temperature variations and juvenile Atlantic salmon in the uppermost areas of the river, experiencing the lowest ambient temperatures during the growth period, displayed the highest growth rates. Within the baselines set by temperature, biotic and abiotic factors connected to water flow regime and variation in food availability are suggested to be a major determinants of the temporal and spatial variation in juvenile Atlantic salmon growth rates.     

Lacustrine growth of juvenile pink salmon and a comparison with sympatric sockeye salmon

The growth rates of naturally sympatric juvenile pink Oncorhynchus gorbuscha and sockeye Oncorhynchus nerka salmon were compared in a common lacustrine environment in south‐west Alsaka, an unusual opportunity given the normal disparity in freshwater residence time of these two species. Fork length ( L _F) frequency distributions of juvenile pink salmon caught in the lake during the summer in 1991 and 1999–2003 indicated a growth rate of 0·54 mm day⁻¹, 54% greater than the estimated growth rate of juvenile sockeye salmon sampled from 1958 to 2003 (0·35 mm day⁻¹). Examination of daily growth rings on otoliths indicated that pink salmon in Lake Aleknagik grew an average of 1·34 mm day⁻¹ in 2003 but sockeye salmon grew only 0·63 mm day⁻¹(average specific growth rates were 3·0 and 1·8% day⁻¹, respectively). Pink salmon increased from c . 32 mm L _F and 0·2 g at emergence to 78 mm L _F and 3·0 g within 3–4 weeks. After experiencing these rapid growth rates, the pink salmon appeared to leave the lake by late July in most years. The diets of pink and sockeye salmon in the littoral zone of the lake were very similar; >80% of the stomach contents consisted of adult and pupal insects and the remainder was zooplankton. This high degree of diet overlap suggested that the observed differences in growth rate were not attributable to variation in prey composition.     

Effects of a founder event and supplementary introductions on genetic variation in a captive breeding population of the endangered Spanish killifish

Twelve polymorphic allozyme loci were employed to assess the genetic change in a captive breeding population of the endangered killifish Aphanius baeticus in the Doñana National Park, south‐western Spain. The initial founder event did not significantly reduce the allelic richness or the expected heterozygosity. No genetic bottleneck signature was detected by tests for deviation from mutation‐drift equilibrium. The F _ST between the wild source and captive population, however, was relatively high (0·053 or 0·122 when excluding or including the locus IDHP‐1 * respectively), after just two to three generations in captivity. Two generations after the incorporation of 68 new wild specimens (greater than five generations after founding) decreased the genetic differences and the F _ST(0·041 excluding IDHP‐1 *). The restoration efforts appeared to be helpful and the study of 12 polymorphic loci and a sensitive parameter such as F _ST were useful for monitoring genetic changes in captivity. Nonetheless, future monitoring should include additional highly polymorphic loci (microsatellites) to achieve higher power to detect genetic change. Such restoration and monitoring efforts should help to avoid rapid inbreeding, adaptation to captivity, and to maintain the long‐term evolutionary potential in small isolated populations.     

A preliminary investigation of allozyme genetic variation and population geographical structure in Aphanius fasciatus from Italian brackish-water habitats

A total of 150 individuals from Aphanius fasciatus from coastal brackish-water habitats was analysed by allozyme electrophoresis to collect data on its genetic variation. From 22 enzymes, 43 putative enzyme-coding loci were resolved, 12 of which were polymorphic at P_0·99 level. Only one of the 31 probability tests showed a significant departure from the Hardy-Weinberg equilibrium. Aphanius fasciatus showed low levels of genetic polymorphism, with expected heterozygosity values ranging from 0·027 (S.E.=0·013) to 0·064 ( s.e. =0·023). Nei's genetic distances between populations ranged from 0·002 to 0·042. Weir & Cockerham F -statistics showed high levels of genetic heterogeneity among populations (jackknifed θ=0·302, s.e. =0·045) and estimates of N _m were < 1, indicating restricted gene flow. Significant positive correlation between genetic distance and geographical distance matrices, detected by Mantel's test ( g =1·941; P 0·001), is consistent with the prediction that the species is genetically structured by isolation-by-distance.     

Migration of hatchery‐reared Atlantic salmon and wild anadromous brown trout post‐smolts in a Norwegian fjord system

Hatchery‐reared Atlantic salmon Salmo salar ( n  = 25) and wild anadromous brown trout (sea trout) Salmo trutta ( n  = 15) smolts were tagged with coded acoustic transmitters and released at the mouth of the River Eira on the west coast of Norway. Data logging receivers recorded the fish during their outward migration at 9, 32, 48 and 77 km from the release site. Seventeen Atlantic salmon (68%) and eight sea trout (53%) were recorded after release. Mean migratory speeds between different receiver sites ranged from 0·49 to 1·82 body lengths (total length) per second (bl s⁻¹) for Atlantic salmon and 0·11–2·60 bl s⁻¹ for sea trout. Atlantic salmon were recorded 9, 48 and 77 km from the river mouth on average 28, 65 and 83 h after release, respectively. Sea trout were recorded 9 km from the release site 438 h after release. Only four (23%) sea trout were detected in the outer part of the fjord system, while the rest of the fish seemed to stay in the inner fjord system. The Atlantic salmon stayed for a longer time in the inner part than in the outer parts of the fjord system, but distinct from sea trout, migrated through the whole fjord system into the ocean.     

Physiological impact of sea lice on swimming performance of Atlantic salmon

Atlantic salmon Salmo salar were infected with two levels of sea lice Lepeophtheirus salmonis (0·13 ± 0·02 and 0·02 ± 0·00 sea lice g⁻¹). Once sea lice became adults, the ventral aorta of each fish was fitted with a Doppler cuff to measure cardiac output ( ̇ ), heart rate ( f _H) and stroke volume ( V _S) during swimming. Critical swimming speeds ( U _crit) of fish with higher sea lice numbers [2·1 ± 0·1 BL (body lengths) s⁻¹] were significantly lower ( P  < 0·05) than fish with lower numbers (2·4 ± 0·1 BL s⁻¹) and controls (sham infected, 2·6 ± 0·1 BL s⁻¹). After swimming, chloride levels in fish with higher sea lice numbers (184·4 ± 11·3 mmol l⁻¹) increased significantly (54%) from levels at rest and were significantly higher than fish with fewer lice (142·0 ± 3·7 mmol l⁻¹) or control fish (159·5 ± 3·5 mmol l⁻¹). The f _H of fish with more lice was 9% slower than the other two groups at U _crit. This decrease resulted in ̇ not increasing from resting levels. Sublethal infection by sea lice compromised the overall fitness of Atlantic salmon. The level of sea lice infection used in the present study was lower than has previously been reported to be detrimental to wild Atlantic salmon.     

Evidence for independent origin of two spring‐spawning ciscoes (Salmoniformes: Coregonidae) in Germany

Combined analyses of mitochondrial DNA (mtDNA) and microsatellite loci were performed to assess the genetic differentiation of two spring‐spawning ciscoes from each other and from sympatric Coregonus albula in two German lakes. Polymorphism was screened at six microsatellite loci and mtDNA for a total of 247 and 94 ciscoes, respectively. Microsatellite data showed a weak differentiation between spring‐spawning Coregonus fontanae and sympatric C. albula in Lake Stechlin ( F _ST = 0–0·008), whereas a significant differentiation was observed between spring‐spawning Coregonus lucinensis and sympatric C. albula in Lake Breiter Luzin ( F _ST = 0·013–0·039). A more pronounced genetic difference was observed between both spring‐spawning species ( F _ST = 0·05–0·128). Shared mtDNA haplotypes among sympatric species within both Lake Stechlin and Lake Breiter Luzin were observed, whereas no haplotype was shared between C. fontanae and C. lucinensis . These results suggest an independent origin for spring‐spawning ciscoes in each lake. Evidence is also provided for mtDNA introgression of Coregonus sardinella into C. lucinensis and C. albula in Lake Breiter Luzin. Postglacially, this species or at least a population which showed mtDNA introgression has colonized the Baltic Sea basin up to the glacial margin that was located between Lakes Stechlin and Breiter Luzin.     

Mackerel icefish size and age differences and long‐term change at South Georgia and Shag Rocks

This study analysed the total length ( L _T)‐frequency distribution of mackerel icefish Champsocephalus gunnari at South Georgia and Shag Rocks from nine bottom trawl surveys at South Georgia and eight at Shag Rocks between 1987 and 2002. The estimated mean L _T of age‐classes 1+, 2+, 3+ and 4+ years during January were, respectively, 14·7, 23·5, 29·8 and 35·1 cm at South Georgia. Age‐classes 1+, 2+ and 3+ years were 18·3, 26·2 and 33·8 cm at Shag Rocks. The derived Bertalanffy growth parameters for South Georgia were: L _∞ = 51·7 cm, k  = 0·27 and t ₀ = −0·26. The mean L _T of each age‐class of C. gunnari at Shag Rocks was significantly larger than at South Georgia, equivalent to c . 5 months growth, although the annual growth in L _T was similar. This is further evidence that C. gunnari hatched earlier at Shag Rocks. At South Georgia, the mean L _T of age‐classes 1+ and 3+ years were correlated, and significantly decreased between 1987 and 2002, and were smaller following warmer summers. This decrease in the size of C. gunnari may be the result of reduced food availability linked to climate warming.     

Molecular genetic assessment of bipolarity in the anchovy genus Engraulis

To test three hypotheses accounting for bipolar distributions in Engraulis , seven of eight taxa (except E. eurystole ) were surveyed with allozymes (34 loci) and 521 bp of the mitochondrial DNA cytochrome b gene. Both allozymes and mtDNA sequences revealed deep separations between New World and Old World anchovies with a mean allozyme genetic distance D  = 1·26 and net mtDNA sequence divergence d  = 15%. These values reflected separations of 5 to 10 million years. Contrary to previous phylogenetic hypotheses, which place north‐east Pacific E. mordax and south‐east Pacific E. ringens as sister taxa, the south‐west Atlantic E. anchoita and E. ringens are most closely related to each other. The north‐east Pacific E. mordax is the closest lineage to the Old World Engraulis , a group of taxa showing low divergences typical of population‐level separations (mean D  = 0·06; mean d  = 0·87%). Bipolarities of sister taxa in the east Atlantic and west Pacific reflect recent dispersals. Bipolarities in the east Pacific and west Atlantic represent paraphyletic taxa in lineages isolated since the Miocene. None of the bipolarities can be attributed to tectonic separations or competitive displacements from the tropics, but the latter situation should be re‐evaluated with comparisons to tropical anchovies of the New World.     

Microsatellite loci reveal highly significant genetic differentiation among Atlantic salmon (Salmo salar L.) stocks from the east coast of Canada

Allele frequency data from eight microsatellite loci provide evidence of highly significant genetic differentiation among stocks of Atlantic salmon Salmo salar L. from the Bay of Fundy, eastern and north-western Nova Scotia and Newfoundland. Estimates of genetic structure ( R _ST and θ) were significant both among all samples taken from the different geographical locations and among samples from geographical regions for which more than one stock was sampled. Samples from the Bay of Fundy taken from stocks which are phenotypically and behaviourally diverse showed particularly high levels of genetic structure. Rogers', allele sharing and (δμ)² distances also revealed significant differences among stock samples and were significantly correlated [Rogers' and (δμ)²] with sea distance between rivers. Results suggest that stocks of Atlantic salmon in eastern Canada are highly diverse genetically and that this should be an important consideration in any management programme for stocks in the area.     

Growth variation in larval Makaira nigricans

The Atlantic blue marlin Makaira nigricans larvae were collected from Exuma Sound, Bahamas and the Straits of Florida over three summers (2000–2002). Sagittal otoliths were extracted and read under light microscopy to determine relationships between standard length ( L _S) and age for larvae from each year and location. Otolith growth trajectories were significantly different between locations: after the first 5–6 days of life, larvae from Exuma Sound grew significantly faster than larvae from the Straits of Florida. Exponential regression coefficients were similar among years for Exuma Sound larvae (mean instantaneous growth rate, G _L = 0·125), but differed between years for larvae from the Straits of Florida ( G _L = 0·086–0·089). Differences in larval growth rates between locations resulted in a 4–6 mm difference in L _S by day 15 of larval life. These differences in growth appeared to be unrelated to mean ambient water temperatures, and may have been caused by location‐specific differences in prey composition or availability. Alternatively, population‐specific differences in maternal condition may have contributed to these differences in early larval growth.     

Ontogenetic changes in temperature preference of Atlantic cod

Final thermal preferendum ( T ) experiments were conducted in a horizontal thermal gradient tank from the beginning of August 2001 to mid‐November 2001 using Atlantic cod Gadus morhua from 6·5 to 79·0 cm fork length ( L _F). The value of T varied significantly ( P  < 0·005) with L _F( T  = 7·23–0·054 L _F), with smaller (younger) fish choosing higher temperatures than larger (older) fish. The preferendum varied from 6·9° C for fish of 6·5 cm to 3·0° C for those of 79·0 cm. Experiments comparing fish positions in the gradient tank between thermal gradients of 0·5–11·0 and 4·5–14·5° C demonstrated that fish positions were determined by temperature selection instead of undesirable tank effects. This study is the first to demonstrate the effect of ontogeny on temperature preferences of a marine fish species.     

The effect of natural selection on estimates of genetic divergence among populations of the Atlantic salmon

The effect of natural selection on the _mMEP-2 * locus on measures of genetic divergence among Atlantic salmon populations was investigated by examining the pattern of change in the level of genetic differentiation (F_ST) averaged over loci when data on the _mMEP-2 * locus were either included or excluded. The level of F_ST among populations at various geographic scales was estimated from allele frequencies at up to four loci (_s AAT-4 *, IDDH-1 *, IDHP-3 *, and _mMEP-2 *). At smaller geographic scales (within river systems or limited geographic regions) levels of variance in _mMEP-2 * allele frequencies were reduced relative to mean levels. At larger geographic scales (across continents or the species range) variation in _mMEP-2 * allele frequencies was greater than mean levels. These results suggest an a priori hypothesis for the effect of selection on the _mMEP-2 * locus which may be applied in future studies on variation in protein coding or other (e.g. mini- and microsatellite) loci in the Atlantic salmon. It is recommended that estimates of gene flow among populations of the Atlantic salmon based on mean F _ST estimates which include data on the _mMEP-2 * locus should be viewed with caution.     

Microsatellite variation in cod Gadus morhua throughout its geographic range

Previous genetic studies using neutral markers such as allozymes, mtDNA and minisatellite loci have demonstrated varying amounts of population structure in cod Gadus morhua throughout the Atlantic. Microsatellite loci, which are potentially the most informative of presently available neutral genetic markers, have been applied extensively within western and eastern Atlantic areas but not on a range-wide basis. In the present study, six microsatellite DNA loci were used to screen cod samples from nine locations throughout the geographic range from the Scotian Shelf in the West Atlantic to the Barents and Baltic Seas in the east. Overall F _ST value was 0·03 ( P = < 0·001) across all samples. Statistically significant population differences over all loci combined were evident between more geographically distant samples, using either heterogeneity tests or F _ST analysis, with at least one locus showing significant differences between all samples (prior to Bonferroni correction). A significant correlation was observed between genetic and geographical distance, suggesting a higher level of historical and contemporary gene flow between adjacent populations than more distant populations. Samples from either end of the geographic range (Scotian Shelf and Baltic Sea) were particularly distinct when analysed using the STRUCTURE programme and also showed a high level of self-assignment when individuals of either the Scotian Shelf or Baltic Sea were tested against the entire data set. The present microsatellite study demonstrates a high level of geographic population structure between the western Atlantic, middle and eastern Atlantic and Baltic Sea, and thus, the findings should be useful in devising overall management and conservation strategies for the species.     

Short‐term freshwater exposure benefits sea lice‐infected Atlantic salmon

Atlantic salmon Salmo salar were infected with sea lice Lepeophtheirus salmonis (0·08 ± 0·007 sea lice g⁻¹) over a period of 4 h. Both infected and non‐infected fish were swim tested in sea water (SW) and fresh water (FW). The ventral aorta of each fish was fitted with a Doppler cuff in order to measure cardiac output, stroke volume and heart rate during swim testing. Blood samples were taken at rest and after exercise. Critical swimming speed of infected fish in SW (2·14 ± 0·08 body lengths, bl s⁻¹) was significantly lower ( P  < 0·05) than infected fish switched to FW (2·81 ± 0·08 bl s⁻¹) and non‐infected fish in SW (2·42 ± 0·04 bl s⁻¹) and FW (2·61 ± 0·08 bl s⁻¹). Cardiac and blood results indicated infected fish exposed to FW did experience stress, but relief from osmotic and ionic distress probably reduces energy expenditure, allowing the increase in performance. As the performance of sea lice‐infected fish improved upon transfer to FW, it is likely that heavily infected salmonids do return to FW to restore compromised osmotic and ionic balance, and remove sea lice in the process.     

The effect of temperature and acclimation period on repeat swimming performance in cutthroat trout

Hatchery cutthroat trout Oncorhynchus clarki clarki were used to examine the effects of 48 h and 3 week temperature acclimation periods on critical swimming speed ( U _crit). The U _crit was determined for fish at acclimation temperatures of 7, 14 and 18° C using two consecutive ramp‐ U _crit tests in mobile Brett‐type swim tunnels. An additional group was tested at the stock's ambient rearing temperature of 10° C. The length of the temperature acclimation period had no significant effect on either the first or the second U _crit( U _crit‐1 and U _crit‐2, respectively) or on the recovery ratio (the quotient of U _crit‐2  U _crit‐1⁻¹). As anticipated, there was a significant positive relationship between U _crit‐1 and temperature ( P  < 0·01) for both acclimation periods, and an increasing, though non‐significant, trend between U _crit‐2 and temperature ( P  = 0·10). Acclimation temperature had no significant effect ( P  = 0·71) on the recovery ratio. These results indicate that a 48 h acclimation to experimental temperatures within the range of −3 to +8° C of the acclimation temperature may be sufficient in studies of swimming performance with this species. This ability to acclimate rapidly is probably adaptive for cutthroat trout and other species that occupy thermally variable environments.