Genetic structure of freshwater Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) populations from the lakes Onega and Ladoga of northwest Russia and implications for conservation

Freshwater Atlantic salmon (Salmo salar L.) populations are a living example of adaptation to the changing conditions caused by glacial cycles. The uniqueness of these populations is emphasized by almost complete resistance to the dangerous parasite Gyrodactylus salaris. In Europe, freshwater salmon populations occur primarily in north-western Russia in the republic of Karelia. These systems include Lakes Ladoga and Onega, the two largest lakes in Europe, each of which harbours a number of freshwater salmon spawning rivers. We used microsatellite markers to study the genetic structure and temporal stability of 11 freshwater salmon populations in Russian Karelia. Populations clustered according their region of origin. Although temporal variation in allele frequencies was observed in the majority of temporal comparisons, various lines of evidence demonstrated that this influence was relatively minor compared to spatial variation that explained eight times more of the variability than temporal variation. Temporal stability tended to occur in populations from rivers with a higher linear lake coefficient. The high level of genetic structuring observed in both lake systems and the apparent low level of migration between populations suggests that treating each river as a separate management unit is recommended. In addition, as the number of populations is large, the best strategy for such fine scale management would be to ensure that the level of natural reproduction in each river is sufficient to sustain the population. A prioritization strategy for population conservation based on estimating the relative roles of different evolutionary forces shaping the gene pools highlighted a number of populations where further monitoring is warranted and also identified populations which could be prioritized for conservation as living gene banks in the event that conservation resources are limited. This prioritization agreed well with the occurrence of temporal (in)stability.     

Long-term temporal changes of genetic composition in brown trout (Salmo trutta L.) populations inhabiting an unstable environment

The genetic structure of brown trout (Salmo trutta) populations inhabiting rivers on the island of Bornholm in the Baltic Sea was studied on a spatial and temporal scale. Low water levels in the rivers during the summer period are assumed to have a significant impact on the persistence of local populations, possibly resulting in a metapopulation structure. Extinctions may, however, also be buffered by a remnant strategy, whereby juveniles escape to river outlets during periods of drought. We compared polymorphism at seven microsatellite DNA loci in contemporary and past samples collected from 1944 to 1997. A principal component analysis, a hierarchical gene diversity analysis and assignment tests showed that the genetic composition of populations was not temporally stable, and that temporal genetic differentiation was much stronger than spatial differentiation. Genetic variability was high and stable over time. Effective population sizes (Ne) and migration rate (m) were estimated using a maximum-likelihood-based implementation of the temporal method. Ne estimates were low (ranging from 8.3 to 22.9) and estimates of m were high (between 0.23 and 0.99), in contrast to other Danish trout populations inhabiting larger and more environmentally stable rivers (Ne between 39.2 and 289.9 and m between 0.01 and 0.09). Thus, the observed spatio-temporal patterns of genetic differentiation can be explained by drift in small persisting populations, where levels of genetic variation are maintained by strong gene flow. However, observations of rivers devoid of trout suggested that population turnover also takes place. We suggest that Bornholm trout represent a metapopulation where the genetic structure primarily reflects strong drift and gene flow, combined with occasional extinction-recolonization events.     

Stocking‐related patterns of genetic variation at enzymatic loci in south European Atlantic salmon populations

Genetic variation at 24 enzymatic loci was investigated in eight south European Atlantic salmon populations from the rivers Nivelle, Cares, Sella, Narcea, Esva, Navia, Porcia and Eo. In these rivers, management based on supplementation of native populations with foreign stocks was carried out for more than one decade. Population genetic patterns expressed in terms of allele frequencies, mean heterozygosity and conformity to Hardy‐Weinberg equilibrium, were significantly different between populations. Relevant temporal changes of genetic variability were reported. Evidence that foreign stocking has disturbed the genetic patterns of some of the studied populations is presented.     

Genetic diversity of north Okhotsk Sea chum salmon populations with natural and artificial reproduction

The variability of 32 enzyme loci was studied in chum salmon populations with different types of reproduction—natural, mixed, and artificial—in some Magadan Region rivers. Among the populations studied, the values of mean heterozygosity and allele number per locus did not differ significantly. We found evidence of definite temporal stability of the populations, and also found that their genetic variability was expressed only slightly but still remained in spite of periodic egg transplantations between rivers. Statistically significant spatial genetic differentiation of the populations accounted for 0.55 to 0.76% of the total variation and the mean inter-year differentiation accounted for 0.30% of the total. Significant temporal (seasonal) genetic subdivision was revealed in chum salmon of the Tauy River. The populations of the Okhotsk Sea coast are very similar genetically to the east Sakhalin populations. The industrial chum salmon population founded and reproduced artificially in the Kulkuty River preserves the genetic similarity of the donor Yama River chum salmon. In the industrial population, we observed a tendency toward reduction of genetic variation over time. The contribution of the Yama population to the gene pool of the Ola chum salmon, (both by natural reproduction and by farming) is small in spite of many large-scale transplantations. However, the consequences of those transplantations are revealed by means of linkage disequilibrium analysis.     

Fitness and sexual response to population density in Daphnia pulex

1. The switch between asexual and sexual reproduction is an important fitness component in cyclically parthenogenetic populations as it is the key to persistence in unstable habitats and because it influences population genetic characteristics such as linkage disequilibrium and population genetic structure.
2. Genetic variation for sexual and asexual reproductive rate ( R _C) was examined, under varying population density, in Daphnia pulex sampled from a rockpool system and two other distinct European localities.
3. Density affected the switch to sexual behaviour, as found in other studies and reduced R _c. Production of males was not correlated with sexual reproduction, promoting outcrossing.
4. Genetic variation was found for the response to density, both within and between populations. There was substantial variation within the rockpools despite their demographic instability, indicating rapid recovery of life-history variation or maintenance during bottlenecks.     

Assessing temporal and spatial variation in wild populations of Atlantic salmon with particular reference to Asturias (Northern Spain) rivers

The extent of genetic variation and levels of temporal and spatial heterogeneity was investigated, at six polymorphic protein‐coding loci, in wild Atlantic salmon Salmo salar populations from six rivers of Asturias (Northern Spain). Also, stocks from northern Europe that were among those introduced to repopulate Asturian Rivers, and other wild Spanish and European populations were characterized. The lack of temporal variation observed suggests that effective population sizes of Asturian populations are sufficiently large to prevent extreme levels of genetic drift and that the introduced fish had a negligible contribution to the fisheries of Asturian rivers.     

The architecture of river networks can drive the evolutionary dynamics of aquatic populations

It is widely recognized that physical landscapes can shape genetic variation within and between populations. However, it is not well understood how riverscapes, with their complex architectures, affect patterns of neutral genetic diversity. Using a spatially explicit agent‐based modeling (ABM) approach, we evaluate the genetic consequences of dendritic river shapes on local population structure. We disentangle the relative contribution of specific river properties to observed patterns of genetic variation by evaluating how different branching architectures and downstream flow regimes affect the genetic structure of populations situated within river networks. Irrespective of the river length, our results illustrate that the extent of river branching, confluence position, and levels of asymmetric downstream migration dictate patterns of genetic variation in riverine populations. Comparisons between simple and highly branched rivers show a 20‐fold increase in the overall genetic diversity and a sevenfold increase in the genetic differentiation between local populations. Given that most rivers have complex architectures, these results highlight the importance of incorporating riverscape information into evolutionary models of aquatic species and could help explain why riverine fishes represent a disproportionately large amount of global vertebrate diversity per unit of habitable area.     

Population genetics of the aquatic fungus Tetracladium marchalianum over space and time

Aquatic hyphomycete fungi are fundamental mediators of energy flow and nutrient spiraling in rivers. These microscopic fungi are primarily dispersed in river currents, undergo substantial annual fluctuations in abundance, and reproduce either predominantly or exclusively asexually. These aspects of aquatic hyphomycete biology are expected to influence levels and distributions of genetic diversity over both spatial and temporal scales. In this study, we investigated the spatiotemporal distribution of genotypic diversity in the representative aquatic hyphomycete Tetracladium marchalianum. We sampled populations of this fungus from seven sites, three sites each in two rivers in Illinois, USA, and one site in a Wisconsin river, USA, and repeatedly sampled one population over two years to track population genetic parameters through two seasonal cycles. The resulting fungal isolates (N = 391) were genotyped at eight polymorphic microsatellite loci. In spite of seasonal reductions in the abundance of this species, genotypic diversity was consistently very high and allele frequencies remarkably stable over time. Likewise, genotypic diversity was very high at all sites. Genetic differentiation was only observed between the most distant rivers (∼450 km). Clear evidence that T. marchalianum reproduces sexually in nature was not observed. Additionally, we used phylogenetic analysis of partial β-tubulin gene sequences to confirm that the fungal isolates studied here represent a single species. These results suggest that populations of T. marchalianum may be very large and highly connected at local scales. We speculate that large population sizes and colonization of alternate substrates in both terrestrial and aquatic environments may effectively buffer the aquatic populations from in-stream population fluctuations and facilitate stability in allele frequencies over time. These data also suggest that overland dispersal is more important for structuring populations of T. marchalianum over geographic scales than expected.     

Lacustrine spatial distribution of landlocked Atlantic salmon populations assessed across generations by multilocus individual assignment and mixed-stock analyses

The objective of this study was to assess the spatiotemporal distribution of four landlocked Atlantic salmon (Salmo salar) populations during their sympatric feeding phase in lake St-Jean (Québec, Canada). A total of 1100 fish captured over a period of 25 years was genotyped at six microsatellite loci in order to assess the temporal stability of the relative proportion of each population in different lake sectors using both individual-based assignment and mixed-stock analysis. Estimates of relative proportions obtained from both methods were highly correlated. A nonrandom spatial distribution of populations was observed for each period and, despite the fact that the overall proportion of each population varied over time, the pattern of differential distribution remained generally stable over time. Furthermore, there were indications that the extent of horizontal spatial overlap among populations was negatively correlated with that of their genetic differentiation at both microsatellites and a major histocompatibility complex locus, and independent of the geographical distance between the rivers of origin. We discuss the hypothesis that the temporal stability of spatial distribution, the lack of an association between spatial partitioning and geographical distance between rivers of origin, and the apparent negative correlation between spatial overlap and genetic differentiation, reflect the outcome of selective pressures driving behavioural differences for spatial niche partitioning among populations.     

Temporal change in genetic structure and effective population size in steelhead trout (Oncorhynchus mykiss)

There is a wealth of published molecular population genetic studies, however, most do not include historic samples and thus implicitly assume temporal genetic stability. We tested for changes in genetic diversity and structure in three populations of steelhead trout (Oncorhynchus mykiss) from a northern British Columbia watershed using seven microsatellite loci over 40 years. We found little change in genetic diversity (mean allele numbers and observed and expected heterozygosity), despite large variation in the estimated numbers of steelhead returning to the watershed over the same time period. However, the temporal stability in genetic diversity is not reflected in population structure, which appears to be high among populations, yet significantly variable over time. The neighbour-joining tree showed that, overall, two of the populations (Zymoetz and Kispiox) clustered separately from the third (Babine); a finding which was not consistent with their geographical separation. The clustering pattern was also not temporally consistent. We used the temporal method to estimate the effective number of breeders (Nb ) for the three populations; our values (Nb = 17-102) were low for the large and presumed vigorous populations of steelhead trout sampled. The low Nb values were also not consistent with the generally high genetic diversity estimates, suggesting the possibility of intermittent gene flow among the three populations. The use of temporal analyses in population genetic samples should be a priority; first, to verify observed patterns in contemporary data, and second, to build a dataset of temporal analyses to allow generalizations to be made concerning temporal genetic stability and effective population size in natural populations.     

Genetic differentiation among populations of Caridinazebra (Decapoda: Atyidae) in tropical rainforest streams, northern Australia

1. Caridina zebra is a common atyid shrimp in some tropical rainforest streams in far north Queensland, Australia. Genetic variation at five allozyme loci was used to estimate the level of dispersal among populations of this species, within and between stream systems. Shrimps were sampled from nine streams in the Tully River catchment and two headwater streams in the adjacent Herbert River catchment in an area under consideration for extensive hydroelectric development.
2. High levels of genetic differentiation were recorded among most populations which suggests that, like other fully aquatic species, movement is limited to a very small spatial scale.
3. In the Tully catchment, populations of shrimp from streams with confluences at high altitude showed less genetic differentiation than those from streams which directly entered the lower river. Dispersal between the latter streams is clearly limited by the presence of large waterfalls and cascades.
4. Adjacent stream populations were often highly differentiated, despite their close proximity, suggesting that overland dispersal is unlikely. However, populations of shrimp in the two streams in the Herbert catchment were strikingly similar in genetic structure to those in adjacent headwater streams of the Tully. Such similarity may reflect relatively recent changes in drainage patterns.     

Population genetic diversity of Prenant’s schizothoracin, Schizothorax prenanti, inferred from the mitochondrial DNA control region

Wild populations of Prenant’s schizothoracin, Schizothorax prenanti, were collected from the Qingyi, Dadu, and Muli rivers in upper reaches of the Yangtze River. Genetic variation and population structure were examined based on analysis of the mitochondrial DNA control region. Nucleotide sequence analysis defined 30 haplotypes in 41 individuals. The nucleotide and haplotype diversities were very high, and genetic distances between the Dadu and Qingyi populations were much smaller than between either of them and the Muli population. The analysis of molecular variance demonstrated that most variation occurred within samples, and the differentiation between Muli and Dadu or Qingyi was significant. Estimates of gene flow indicated reproductive isolation between Muli and the other populations, and this divergence might be related to geographical location. The results are mostly consistent with the findings from AFLP analysis and also suggested considerable genetic diversity of the populations in the Muli, Dadu, and Qingyi rivers.     

The effect of dispersal ability in winter and summer stoneflies on their genetic differentiation

Abstract.  1. Plecopteran species disperse less than most other aquatic insects. Within stoneflies, members of different families vary in the degree of wing morphology and season of adult emergence.
2. Dispersal limitations were tested to determine if there were increased differences among the nearby, Chagrin and Grand River populations in north-eastern Ohio, by comparing genetic variation within the 16s rRNA region of mitochondrial DNA in two stoneflies. Allocapnia recta emerges in winter and often has rudimentary wings, and Leuctra tenuis emerges in summer with fully developed wings.
3. There was significant genetic variability between the samples of A. recta from two adjacent rivers (FST = 0.20), but not between samples of L. tenuis (FST = 0.07).
4. Distinct clades in A. recta were found to occur within the minimum spanning tree specific to the Chagrin River, which contributed to a significant difference in gene diversity between the two rivers. Haplotypes in L. tenuis appeared randomly distributed between the two rivers; however, nucleotide diversity was significantly less in samples from the Grand River.
5. Shared haplotypes of both species illustrate the inter-connectedness of the Chagrin and Grand River populations, and the lower genetic variability of L. tenuis between the two rivers is indicative of its greater dispersal capability.     

汩罗江与柳江鳤鱼线粒体细胞色素b基因的遗传变异初探

测定并分析了长江水系汨罗江4尾鳤鱼(Ochetobius elongates)和珠江水系柳江2尾鳤鱼的线粒体细胞色素b基因部分序列981bp,发现6个单倍型,其中19个变异位点,简约信息位点15个;AT含量和GC含量分别为57.6%和42.4%。在基于Kimura双参数模型的邻接树上,汩罗江和柳江鳤鱼各自聚群。汨罗江4尾鳤鱼间的遗传距离为0.001～0.007,柳江2尾鳤鱼间遗传距离为0.001,而2群体间的平均遗传距离为0.015(0.013～0.017),群体间的遗传变异大于群体内的遗传变异,表明汩罗江鳤鱼与柳江鳤鱼出现较明显的遗传分化。由于本研究采集地理群体少,分析的样本小,为更好地揭示长江和珠江水系鳤鱼的遗传变异,上述结论尚需进一步分析更多大样本的地理群体加以验证。     

Gene flow between populations of two invertebrates in springs

1. Using allozymes, we analysed genetic structure of the freshwater gastropod Bythinella dunkeri and the freshwater flatworm Crenobia alpina. The two species are habitat specialists, living almost exclusively in springs. The sampled area in Hesse (Germany) covers a spatial scale of 20 km and includes two river drainages. From the biology of the two species we expected little dispersal along rivers. However, the possibility exists that groundwater provide suitable pathways for dispersal. 2. In B. dunkeri heterozygosity decreased from west to east. For some alleles we found clines in this geographic direction. These clines generated a positive correlation between geographic distance and genetic differentiation. Furthermore patterns of genetic variation within populations suggested that populations may have been faced with bottlenecks and founder effects. If populations are not in population genetic equilibrium, such founder effects would also explain the rather high amount of genetic differentiation between populations (10%). 3. For C. alpina the mean number of alleles decreased with increasing isolation of populations. Genetic differentiation between populations contributed 19% to the total genetic variation. Genetic differentiation was not correlated to geographic distance, but compared with B. dunkeri variability of pairwise differentiation between pairs of populations was higher in C. alpina. 4. Overall B. dunkeri appears to be a fairly good disperser, which may use groundwater as dispersal pathway. Furthermore populations seem to be not in equilibrium. In contrast C. alpina forms rather isolated populations with little dispersal between springs and groundwater seems to play no important role for dispersal.     

Allozyme variability in brown trout Salmo trutta in Chile

1. Brown trout ova were imported during the last century from different locations in Europe to establish populations in Chilean rivers (South America). The rivers are currently occupied by naturalized populations that have adapted to very different environmental conditions, such as areas of semi-desert in the north, or rainy and cold areas in the south.
2. In this first study in this geographical area, electrophoretic variability of proteins encoded by twenty-five loci was screened in seven populations from northern to southern Chile.
3. The results show significant heterogeneity of allelic frequencies between populations in seven of eleven polymorphic loci detected. The estimated value of genetic diversity 0.1274 ( H _T) is higher than that observed in populations from areas of natural distribution of this species. However, only 12.64% of this genetic diversity was found between samples ( G _ST), indicating a low genetic divergence among Chilean populations. The observed associations among the Chilean and 'modern' group of European populations suggests the probable origin of the new populations.     

The genetic underpinnings of population cyclicity: establishing expectations for the genetic anatomy of cycling populations

Despite extensive research into the mechanisms underlying population cyclicity, we have little understanding of the impacts of numerical fluctuations on the genetic variation of cycling populations. Thus, the potential implications of natural and anthropogenically‐driven variation in population cycle dynamics on the diversity and evolutionary potential of cyclic populations is unclear. Here, we use Canada lynx Lynx canadensis matrix population models, set up in a linear stepping‐stone, to generate demographic replicates of biologically realistic cycling populations. Overall, increasing cycle amplitude predictably reduced genetic diversity and increased genetic differentiation, with cyclic effects increased by population synchrony. Modest dispersal rates (1–3% of the population) between high and low amplitude cyclic populations did not diminish these effects suggesting that spatial variation in cyclic amplitude should be reflected in patterns of genetic diversity and differentiation at these rates. At high dispersal rates (6%) groups containing only high amplitude cyclic populations had higher diversity and lower differentiation than those mixed with low amplitude cyclic populations. Negative density‐dependent dispersal did not impact genetic diversity, but did homogenize populations by reducing differentiation and patterns of isolation by distance. Surprisingly, temporal changes in diversity and differentiation throughout a cycle were not always consistent with population size. In particular, negative density‐dependent dispersal simultaneously decreased differences in genetic diversity while increasing differences in genetic differentiation between numerical peaks and nadirs. Combined, our findings suggest demographic changes at fine temporal scales can impact genetic variation of interacting populations and provide testable predictions relating population cyclicty to genetic variation. Further, our results suggest that including realistic demographic and dispersal parameters in population genetic models and using information from temporal changes in genetic variation could help to discern complex demographic scenarios and illuminate population dynamics at fine temporal scales.     

Dispersal and recruitment of Tasiagma ciliata (Trichoptera: Tasimiidae) in rainforest streams, south-eastern Australia

1. This study examined genetic variation within and among populations of the caddis fly Tasiagma ciliata (Tasimiidae: Trichoptera) from rainforest streams in south-east Queensland, Australia.
2. Very low levels of genetic differentiation at large spatial scales, between subcatchments and between catchments, indicated that dispersal by the winged adults is widespread. However, significant genetic differentiation at the smallest spatial scale examined, within reaches in a single stream, suggested limited movement by larvae within streams.
3. A patchy distribution of deviations from Hardy–Weinberg equilibrium and differences in patterns among allozyme loci suggested that populations in particular reaches were the result of only a few matings.
4. These results are surprising, given the large numbers of larvae present within a single reach. We suggest that stochastic effects of recruitment may underlie much of the spatial and temporal variation in population numbers in these rainforest streams.     

Fine-scale genetic structure of Atlantic salmon (Salmo salar) using microsatellite markers: effects of restocking and natural recolonization

1. An important goal of conservation biology is to preserve the evolutionary potential of a species by maintaining natural levels of genetic diversity. Here, we assess the population differentiation in the Atlantic salmon, Salmo salar, listed in Annex II of the European Habitats Directive, to provide valuable information for its conservation in Normandy (France).
2. Samples collected from 10 natural sites revealed that 13 of 14 microsatellite loci were polymorphic. Significant differentiation among populations was detected ( F _ST = 0.054, P  <   0.001), and all F _ST pairwise comparisons except one were significant. A genetic split was observed between populations inhabiting streams with limestone geology compared to those inhabiting streams with siliceous geology, which could reflect adaptative differences.
3. Hatchery stocks used for the restocking of two rivers were genetically distinct from native stocks.
4. Analysis of three stream habitats restored in 1995 showed that all were recolonized naturally by wild salmon from geographically close populations and no founder effects were detected. Allelic richness was similar between recolonized and wild populations.
5. From a management perspective, our study revealed that restoration of habitat is very effective to recreate new populations in rivers from which salmon have disappeared and that natural recolonization can be fast and effective in terms of genetic diversity.