Per Erik Jorde

Fine-scaled geographical population structuring in a highly mobile marine species: the Atlantic cod.

Compared with many terrestrial and freshwater environments, dispersal and interbreeding is generally much less restricted in the marine environment. We studied the tendency for a marine species, the Atlantic cod, to be sub‐structured into genetically differentiated populations on a fine geographical scale. We selected a coastal area free of any obvious physical barriers and restricted sampling to a 300‐km region, well within the dispersal ability of this species. Screening 10 polymorphic microsatellite loci in 6 samples we detected a weak, but consistent, differentiation at all 10 loci. The average FST over loci was small (0.0023) but highly significant statistically, demonstrating that genetically differentiated populations can arise and persist in the absence of physical barriers or great distance. We found no geographical pattern in the genetic differentiation and there was no apparent trend of isolation by distance along the coastline. These findings lend support to the notion that low levels of differentiation are due to passive transport of eggs or larvae by the ocean currents rather than to adult dispersal, the latter being strongly dependent on distance.

Power for detecting genetic divergence: differences between statistical methods and marker loci

Information on statistical power is critical when planning investigations and evaluating empirical data, but actual power estimates are rarely presented in population genetic studies. We used computer simulations to assess and evaluate power when testing for genetic differentiation at multiple loci through combining test statistics or P values obtained by four different statistical approaches, viz. Pearsons chi‐square, the log‐likelihood ratio G‐test, Fishers exact test, and an FST‐based permutation test. Factors considered in the comparisons include the number of samples, their size, and the number and type of genetic marker loci. It is shown that power for detecting divergence may be substantial for frequently used sample sizes and sets of markers, also at quite low levels of differentiation. The choice of statistical method may be critical, though. For multi‐allelic loci such as microsatellites, combining exact P values using Fishers method is robust and generally provides a high resolving power. In contrast, for few‐allele loci (e.g. allozymes and single nucleotide polymorphisms) and when making pairwise sample comparisons, this approach may yield a remarkably low power. In such situations chi‐square typically represents a better alternative. The G‐test without Williamss correction frequently tends to provide an unduly high proportion of false significances, and results from this test should be interpreted with great care. Our results are not confined to population genetic analyses but applicable to contingency testing in general.

Unbiased estimator for genetic drift and effective population size.

Amounts of genetic drift and the effective size of populations can be estimated from observed temporal shifts in sample allele frequencies. Bias in this so-called temporal method has been noted in cases of small sample sizes and when allele frequencies are highly skewed. We characterize bias in commonly applied estimators under different sampling plans and propose an alternative estimator for genetic drift and effective size that weights alleles differently. Numerical evaluations of exact probability distributions and computer simulations verify that this new estimator yields unbiased estimates also when based on a modest number of alleles and loci. At the cost of a larger standard deviation, it thus eliminates the bias associated with earlier estimators. The new estimator should be particularly useful for microsatellite loci and panels of SNPs, representing a large number of alleles, many of which will occur at low frequencies.

Are low but statistically significant levels of genetic differentiation in marine fishes ‘biologically meaningful’? A case study of coastal Atlantic cod

A key question in many genetic studies on marine organisms is how to interpret a low but statistically significant level of genetic differentiation. Do such observations reflect a real phenomenon, or are they caused by confounding factors such as unrepresentative sampling or selective forces acting on the marker loci? Further, are low levels of differentiation biologically trivial, or can they represent a meaningful and perhaps important finding? We explored these issues in an empirical study on coastal Atlantic cod, combining temporally replicated genetic samples over a 10‐year period with an extensive capture–mark–recapture study of individual mobility and population size. The genetic analyses revealed a pattern of differentiation between the inner part of the fjord and the open skerries area at the fjord entrance. Overall, genetic differentiation was weak (average FST = 0.0037), but nevertheless highly statistical significant and did not depend on particular loci that could be subject to selection. This spatial component dominated over temporal change, and temporal replicates clustered together throughout the 10‐year period. Consistent with genetic results, the majority of the recaptured fish were found close to the point of release, with <1% of recaptured individuals dispersing between the inner fjord and outer skerries. We conclude that low levels of genetic differentiation in this marine fish can indeed be biologically meaningful, corresponding to separate, temporally persistent, local populations. We estimated the genetically effective sizes (Ne) of the two coastal cod populations to 198 and 542 and found a Ne/N (spawner) ratio of 0.14.

Management implications of genetic differentiation between native and hatchery populations of brown trout (Salmo trutta) in Spain

Abstract Isozyme electrophoresis was used to assess the amount and distribution of genetic variation among natural populations and hatchery stocks of brown trout (Salmo trutta) in Spain. Genetic variation was found at 19 out of 50 protein coding loci. The hatchery stocks were conspicuously different from the natural populations, and the two groups appeared to represent distinctly divergent evolutionary lineages of brown trout. The hatchery stocks were highly polymorphic but also markedly homogeneous; only 3% of the total genetic variation was explained by stock differences, indicating a common origin for these stocks. In sharp contrast to the hatchery stocks, the natural populations were found to be highly divergent, and more than 60% of their total genetic variability was due to differences between populations. There is evidence indicating impending or actual loss of natural populations by intrusions of fish of hatchery origin or ancestry. In the interest of optimal resource management, we recommend that the indigenous populations of Spain be fully identified and protected, and that the existing hatchery stocks be replaced with local natural populations.

Statistical power when testing for genetic differentiation

A variety of statistical procedures are commonly employed when testing for genetic differentiation. In a typical situation two or more samples of individuals have been genotyped at several gene loci by molecular or biochemical means, and in a first step a statistical test for allele frequency homogeneity is performed at each locus separately, using, e.g. the contingency chi‐square test, Fisher’s exact test, or some modification thereof. In a second step the results from the separate tests are combined for evaluation of the joint null hypothesis that there is no allele frequency difference at any locus, corresponding to the important case where the samples would be regarded as drawn from the same statistical and, hence, biological population. Presently, there are two conceptually different strategies in use for testing the joint null hypothesis of no difference at any locus. One approach is based on the summation of chi‐square statistics over loci. Another method is employed by investigators applying the Bonferroni technique (adjusting the P‐value required for rejection to account for the elevated alpha errors when performing multiple tests simultaneously) to test if the heterogeneity observed at any particular locus can be regarded significant when considered separately. Under this approach the joint null hypothesis is rejected if one or more of the component single locus tests is considered significant under the Bonferroni criterion. We used computer simulations to evaluate the statistical power and realized alpha errors of these strategies when evaluating the joint hypothesis after scoring multiple loci. We find that the ‘extended’ Bonferroni approach generally is associated with low statistical power and should not be applied in the current setting. Further, and contrary to what might be expected, we find that ‘exact’ tests typically behave poorly when combined in existing procedures for joint hypothesis testing. Thus, while exact tests are generally to be preferred over approximate ones when testing each particular locus, approximate tests such as the traditional chi‐square seem preferable when addressing the joint hypothesis.

Transport of North Sea cod larvae into the Skagerrak coastal populations

The Atlantic cod (Gadus morhua) is economically one of the worlds most important marine species––a species presently suffering from heavy overexploitation throughout its range of distribution. Although not fully understood, the Atlantic cod is believed to be structured into populations in a rather complex manner, whereby both highly migratory and more confined ocean–spawning stocks coexist with stationary coastal populations. Owing to the complex population structure, little is presently known about how overexploitation of offshore stocks may affect other segments of the species. Here, we use microsatellite DNA analyses of coastal and offshore cod in combination with oceanographic modelling to investigate the population structure of Atlantic cod in the North Sea–Skagerrak area and evaluate the potential for larval transport into coastal populations. Our results suggest an extensive but temporally variable drift of offshore cod larvae into coastal populations. In a year (2001) with high inflow of North Sea waters into the Skagerrak we find that juvenile cod caught along the Skagerrak coast are predominantly of North Sea origin, whereas in a year (2000) with low inflow juveniles appear to be of local origin. These findings indicate that offshore cod may influence coastal cod populations over large distances.

Small‐scale biocomplexity in coastal Atlantic cod supporting a Darwinian perspective on fisheries management

Harvesting of marine resources raises concerns about how to identify and preserve biocomplexity, including the diversity of life histories found within and among wild populations of a species. In order to fully accomplish this, there is a need to elucidate the underlying causes of phenotypic variation, and how this variation responds to environmental changes. In general, both evolutionary (genetic) and nonevolutionary (plastic) responses may occur. Plastic responses to environmental change are expected to shift the phenotype along a reaction norm, while an evolutionary response is expected to shift the reaction norm itself. Here, we assess the maturation patterns of coastal Atlantic cod (Gadus morhua) in Skagerrak, where studies using neutral markers have revealed genetically differentiated populations of this harvested fish within tens of kilometres of coastline. Our results suggest that physiological state prior to the spawning season, as well as juvenile growth, both influence the probability of completing sexual maturation at a given age. Furthermore, our results point towards a spatial structuring of this plasticity (i.e. the maturation reaction norms) comparable with population connectivity inferred from neutral markers. We argue that such fine‐scale biocomplexity calls for a Darwinian approach to fisheries management.

Cryptic population structure in a large, mobile mammalian predator: the Scandinavian lynx

The Eurasian lynx (Lynx lynx) is an example of a species that has gone through a severe bottleneck, leading to near extinction in Scandinavia around 1930 — a pattern shared with several other large carnivorous mammals. Here we extend previous genetic analyses of northern European lynx, confirming that lynx from the Scandinavian Peninsula represent a distinct clade differing clearly from European conspecifics. Furthermore, and despite a recent bottleneck and subsequent range expansion, we detect marked genetic differentiation within Scandinavia. This differentiation is largely manifested as a north–south gradient, with a linear increase in the quantity FST/(1 − FST). Aided by computer simulations we find that this pattern is unlikely to have arisen by random genetic drift in the short time since lynx started to expand in the 1950s, suggesting that the spatial structure may predate the bottleneck. Individual‐based analyses indicate that, instead of a continuous gradient, Scandinavian lynx may be structured into three more or less distinct groups, possibly corresponding to northern, central and southern subpopulations. The presence of such structuring was unknown previously and was unexpected from general considerations on the mobility of the species, historical data and the absence of geographical barriers. Our study demonstrates how molecular markers may be used to detect cryptic population structure, invisible using traditional methods.

Genetic differentiation among populations of the beetle Bolitophagus reticulatus (Coleoptera: Tenebrionidae) in a fragmented and a continuous landscape

The effect of habitat fragmentation on genetic differentiation among local populations of the fungivorous beetle Bolitophagus reticulatus (Coleoptera: Tenebrionidae) was studied in two contrasting landscapes: one heavily fragmented with forest fragments of variable size surrounded by inhabitable agricultural fields, the other an old forest providing a continuous habitat. The genetic structure of the beetle within each of the two contrasting areas was investigated by means of protein electrophoresis, screening four polymorphic loci in 20 populations from each area. In both areas there were significant genetic differences among local populations, but on average differentiation in the fragmented area was three times greater than in the continuous one, strongly indicating a genetic isolation effect of habitat fragmentation. These genetic results are in accordance with previous studies on dispersal in this species.