Islands in the ice: colonisation routes for rock ptarmigan to the Svalbard archipelago

The ancestors of rock ptarmigan Lagopus muta originated in the Beringia area well before the disruption of the Beringian land bridge. They spread westwards to Siberia and eastwards to the North American arctic and Greenland. The distribution of rock ptarmigan has been affected by glaciations restricting it to geographically limited refugia. Today the species has a circumpolar distribution in arctic tundra and alpine habitats, with up to 30 subspecies recognised based on morphological characters. We sequenced the mitochondrial control region for 72 individuals and genotyped 69 individuals for 12 microsatellite loci to investigate neutral genetic variation within and among five rock ptarmigan populations, Greenland, Iceland, Scandinavia, Svalbard and Taimyr. Genetic structure among the studied samples was high, overall F_ST estimated from microsatellite loci was 0.18 and only one out of 16 mtDNA haplotypes was found in more than one population. Genetic variation ( h, π, H_e, allelic richness) was slightly lower in the Svalbard population than in other populations, suggesting a low effective population size, possibly due to isolation following colonisation. An unrooted network and a phylogenetic tree showed that the Scandinavian population has diverged from the other populations by at least ten mutational steps, probably due to independent colonisation of Europe and subsequent long-term isolation, and rules out Scandinavia as a source for colonisation of Svalbard. Alignment with partial control region sequences from other studies showed that the haplotype that was central in our network and found on Svalbard and in Taimyr, most likely corresponds to a haplotype found in Siberia, Alaska and the Canadian Arctic, but not in Greenland, Scandinavia and Iceland. This suggests an eastern origin of rock ptarmigan in Svalbard, although this question cannot be settled conclusively.     

Population structure of Purple Sandpipers (Calidris maritima) as revealed by mitochondrial DNA and microsatellites

The Purple Sandpiper (Calidris maritima) is a medium‐sized shorebird that breeds in the Arctic and winters along northern Atlantic coastlines. Migration routes and affiliations between breeding grounds and wintering grounds are incompletely understood. Some populations appear to be declining, and future management policies for this species will benefit from understanding their migration patterns. This study used two mitochondrial DNA markers and 10 microsatellite loci to analyze current population structure and historical demographic trends. Samples were obtained from breeding locations in Nunavut (Canada), Iceland, and Svalbard (Norway) and from wintering locations along the coast of Maine (USA), Nova Scotia, New Brunswick, and Newfoundland (Canada), and Scotland (UK). Mitochondrial haplotypes displayed low genetic diversity, and a shallow phylogeny indicating recent divergence. With the exception of the two Canadian breeding populations from Nunavut, there was significant genetic differentiation among samples from all breeding locations; however, none of the breeding populations was a monophyletic group. We also found differentiation between both Iceland and Svalbard breeding populations and North American wintering populations. This pattern of divergence is consistent with a previously proposed migratory pathway between Canadian breeding locations and wintering grounds in the United Kingdom, but argues against migration between breeding grounds in Iceland and Svalbard and wintering grounds in North America. Breeding birds from Svalbard also showed a genetic signature intermediate between Canadian breeders and Icelandic breeders. Our results extend current knowledge of Purple Sandpiper population genetic structure and present new information regarding migration routes to wintering grounds in North America.     

Genetic variation and structure of house sparrow populations: is there an island effect?

Population genetic structure and intrapopulation levels of genetic variation have important implications for population dynamics and evolutionary processes. Habitat fragmentation is one of the major threats to biodiversity. It leads to smaller population sizes and reduced gene flow between populations and will thus also affect genetic structure. We use a natural system of island and mainland populations of house sparrows along the coast of Norway to characterize the different population genetic properties of fragmented populations. We genotyped 636 individuals distributed across 14 populations at 15 microsatellite loci. The level of genetic differentiation was estimated using F‐statistics and specially designed Mantel tests were conducted to study the influence of population type (i.e. mainland or island) and geographic distance on the genetic population structure. Furthermore, the effects of population type, population size and latitude on the level of genetic variation within populations were examined. Our results suggest that genetic processes on islands and mainland differed in two important ways. First, the intrapopulation level of genetic variation tended to be lower and the occurrence of population bottlenecks more frequent on islands than the mainland. Second, although the general level of genetic differentiation was low to moderate, it was higher between island populations than between mainland populations. However, differentiation increased in mainland populations somewhat faster with geographical distance. These results suggest that population bottleneck events and genetic drift have been more important in shaping the genetic composition of island populations compared with populations on the mainland. Such knowledge is relevant for a better understanding of evolutionary processes and conservation of threatened populations.     

What can morphology and isozymes tell us about the history of the Dryas integrifolia–octopetala complex?

An earlier study of the morphological variation in the circumpolar genus Dryas in Greenland revealed that populations in east Greenland are highly variable and this variation was interpreted as a hybrid zone between D. octopetala and D. integrifolia. It was proposed that populations of Dryas in east Greenland immigrated postglacially from Svalbard. The purpose of our study was to investigate the population structure of the Dryas integrifolia-octopetala complex using a combination of morphological and genetic characters. Populations of Dryas were sampled in Greenland, Svalbard, Iceland and north Scandinavia (Sweden). The morphological traits clearly separated west Greenland from Svalbard, Iceland and north Scandinavia. A large number of individuals from east Greenland and some from north Greenland could be interpreted as hybrids between the morphs. The genetic pattern was, however, partly different as the greatest amount of differentiation among regions was found between west and north Greenland (FRT = 0.65). The highest degree of genetic variation appeared within east Greenland. This was in accordance with the putative hybrid origin of the populations, as concluded based on the morphological results. The total amount of genetic differentiation found among the six regions (FRT = 0.61) was larger than between the two species (FRT = 0.44). The genetic pattern matched the proposed postglacial migration routes.     

Geographical variation in the breeding biology of the Purple Sandpiper Calidris maritima

Studies in Iceland (66°N) and Svalbard (78°N), combined with the results of previous work, allowed geographical comparisons of different aspects of the breeding biology of the Purple Sandpiper Calidris maritima , testing predictions that they may be related to latitude and altitude (here taken as surrogates for climatic severity). The breeding density was lowest in the polar deserts of Franz Josef Land (82°N) and highest in coastal Arctic Tern Sterna paradisaea colonies in Iceland where the terns perhaps afforded protection from predators. There was no difference in the percentages of 1-year-old (first-year) birds in the breeding populations in Svalbard and Iceland. Time of breeding was related to both latitude and altitude, being later at higher latitudes and altitudes. In Iceland, larger males and males over 1 year old nested earlier than small males and 1-year-old males, respectively. Relaying occurred in Iceland but it was not observed in Svalbard. There were no geographical differences in clutch size. Egg size was related to female size in Svalbard. There were also geographical differences in egg size associated with female size (wing length), with the largest eggs in Iceland and smallest in south Norway. Males usually attended broods and the rates at which females deserted their broods were similar in Iceland and Svalbard. Chick growth rates were similar for Iceland and Franz Josef Land. Thus, in breeding biology, Purple Sandpipers varied geographically only in breeding density, time of nesting and egg size, which was associated with female size.     

Genetic variation in Minuartia (Caryophyllaceae) in Svalbard

Based on a survey of eight enzyme systems, genetic variation in three Minuartia species was analysed and the diversity in 15 populations in Svalbard (78°N) compared to seven populations in Norway, Iceland, and Greenland. In the two sexual diploids, M. biflora and M. rubella , genetic diversity was as high in populations from Svalbard as in populations from more southerly latitudes. In M. biflora , eight out of 15 loci were polymorphic (P=53.3 %); in M. rubella eight out of 18 (P=44.4 %). The mean number of alleles per locus (A) and the number of multilocus genotypes (MG) were also higher in M. biflora than in M. rubella , A=1.60 and MG=21 vs. A=1.44 and MG=9. The proportion of genetic diversity due to variation among populations was much lower in M. biflora than in M. rubella , FST=0.249 vs. FST=0.895, and the estimated gene flow much higher, Nm=0.745 vs. Nm=0.029, indicating that M. biflora is a mixed mater and M. rubella a selfer. In the vegetatively reproducing tetraploid, M. rossii , 13 (65 %) out of 20 putative loci showed fixed heterozygosity, the mean number of alleles per 'locus' was A=1.70, and two multilocus phenotypes were observed, proving that at least two clones occur in Svalbard. No allele was shared by all three species and Nei's genetic identity for biflora-rubella was extremely low, I=0.07. The results suggest that the three species, which are currently placed in separate sections, represent lineages that diverged a long time ago.     

THE MIGRATION PATTERNS OF THE PURPLE SANDPIPER CALIDRIS MARITIMA

Summers, R. W. 1994. The migration patterns of the Purple Sandpiper Calidris maritima. Ostrich 65: 167–173.

The Purple Sandpiper breeds largely in the Arctic, and winters (boreal season) on the rocky shores of the north Atlantic, further north than any other sandpiper. As the populations from Canada, Greenland, Iceland, Svalbard, Norway and Russia differ in wing and bill lengths it is possible to match measurements taken from breeding birds with samples of birds caught in winter. Ringing recoveries, especially from colour marked birds, have also helped to determine migration routes and wintering areas. Four populations move to the nearest ice-free coast. Two populations move south of the nearest ice-free coast, being replaced by larger birds from a more northerly population (“chain migration”). Only the north Canadian population is believed to migrate a long distance, “leap-frogging” other winter populations. These patterns are discussed in relation to theories for the migration patterns of waders.     

Genetic variation at nine short tandem repeat loci among islanders of the eastern Adriatic coast of Croatia

We have analyzed the extent of genetic variation at nine autosomal short tandem repeat loci (D3S1358, VWA, FGA, TH01, TPOX, CSF1PO, D5S818, D13S317, D7S820) among six populations from Croatia: five distributed in the islands of the eastern Adriatic coast and one from the mainland. The purpose is to investigate the usefulness of these loci in detecting regional genetic differentiation in the studied populations. Significant heterogeneity among the island and mainland populations is revealed in the distributions of allele frequencies; however, the absolute magnitude of the coefficient of gene differentiation is small but significant. The summary measures of genetic variation, namely, heterozygosity, number of alleles, and allele size variance, do not indicate reduced genetic variation in the island populations compared to the mainland population. In contrast to the two measures of genetic variation, allele size variance and within-locus heterozygosity, the imbalance index (beta) indicates evidence of recent expansion of population sizes in all islands and in the mainland. High mutation rates of the studied loci together with local drift effects are likely explanations for interisland genetic variation and the observed lack of reduced genetic diversity among the island populations.     

High genetic variability in an ecologically variable vertebrate,Bufo viridis

Allozymic variation in proteins encoded by 26 loci was analyzed electrophoretically in 517 specimens of green toads from 11 populations from Israel and one population from Vis Island in the Adriatic Sea. Genetic variation in this toad is the highest yet reported in any vertebrate. All three genetic parameters, mean number of alleles per locus (A), mean proportion of loci polymorphic per population (P), and mean number of heterozygous loci per individual (H), are very high (A = 1.65, range 1.38-2.04; P = 0.423, range 0.346-0.615; H = 0.133, range 0.108-0.159). Central and marginal mainland populations are only slightly more variable than desert isolates, but much more variable than the Vis Island population. Genetic similarity is very high between mainland populations (S = 0.951, range 0.93-0.97). Frequencies of two alleles (Icd-lc and Tfa) are correlated with an ecological gradient of increasing aridity. Regulatory enzymes appeared to contribute more to overall polymorphism than non-regulatory enzymes. The genetic variation observed suggests that selection for heterozygosity as an adaptive strategy is operating in the ecologically variable environment in which green toads live.     

A population on the edge: genetic diversity and population structure of the world's northernmost harbour seals (Phoca vitulina)

It is crucial to examine the genetic diversity and structure of small, isolated populations, especially those at the edge of their distribution range, because they are vulnerable to stochastic processes if genetic diversity is low and the isolation level is high, and because such populations provide insight into the consequences of population declines in a broader conservation context. The harbour seal (Phoca vitulina) population in Svalbard is the world's northernmost P. vitulina population. Nothing is known about the genetic diversity, distinctiveness, or origin of this small, marginalized mammalian population. Thus, the present study investigated its genetic status in the context of nearby P. vitulina in Iceland, south‐east Greenland, and northern Norway: this species is depleted/threatened in all of these regions. A total of 174 samples distributed between the four locations were analysed using 15 polymorphic microsatellites and variation in the displacement loop region (D‐loop). Each of the four locations was a genetically distinct population. The Svalbard population was highly genetically distinct, had reduced genetic diversity, received limited gene flow, had a rather low effective population size, and showed an indication of having experienced a bottleneck resulting from a recent population decline. The significant heterozygote excess observed in the Svalbard sample might be attributed to the low effective population size, which could initiate future population inbreeding effects. This phenomenon has not been reported earlier from other P. vitulina populations, but if the Svalbard population is experiencing inbreeding, this could reduce its resilience to climate change, disease outbreaks, or other perturbations. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 420–439.     

Reduced MHC class II diversity in island compared to mainland populations of the black-footed rock-wallaby (<Emphasis Type="Italic">Petrogale lateralis lateralis</Emphasis>)

Many animal populations that are endangered in mainland areas exist in stable island populations, which have the potential to act as an “ark” in case of mainland population declines. Previous studies have found neutral genetic variation in such species to be up to an order of magnitude lower in island compared to mainland populations. If low genetic variation is prevalent across fitness-related loci, this would reduce the effectiveness of island populations as a source of individuals to supplement declining mainland populations or re-establish extinct mainland populations. One such species, the black-footed rock-wallaby (Petrogale lateralis lateralis), exists within fragmented mainland populations and small island populations off Western Australia. We examined sequence variation in this species within a fitness-related locus under positive selection, the MHC class II DAB β1 locus. The mainland populations displayed greater levels of allelic diversity (4–7 alleles) than the island population, despite being small and isolated, and contained at least two DAB gene copies. The island population displayed low allelic diversity (2 alleles) and fewer alleles per individual in comparison to mainland populations, and probably possesses only one DAB gene copy. The patterns of DAB diversity suggested that the island population has a markedly lower level of genetic variation than the mainland populations, in concordance with results from microsatellites (genotyped in a previous study), but preserved unique alleles which were not found in mainland populations. Where possible, conservation actions should pool individuals from multiple populations, not only island populations, for translocation programs, and focus on preventing further declines in mainland populations.     

Genetic variation in Arabis petraea, a disjunct species in northern Europe

Variation at 7 isoenzyme loci was studied m 15 populations (6 from Iceland, 4 from W Norway, 4 from central Sweden at the Bothnian Gulf, and 1 from Russian Karelia) Heterozygosity within populations was found to be remarkably high compared with taxa with similar reproductive systems and phytogeographical histories Genetic distances calculated from isoenzyme data showed a close connection between the Norwegian and Swedish populations, indicating a western rather than an eastern immigration route for the isolated Swedish population Leaf shape variation displayed a large scale geographical pattern with strong differences between regions and low variation within populations     

Genetic variation and differentiation in Nordic populations of Elymus alaskanus (Scrib. ex Merr.) Löve (Poaceae)

 To gain information on the extent and nature of genetic variation in Elymus alaskanus, levels and distribution of genetic variation were assessed within and among 13 populations originating from Iceland, Norway, Sweden and Russia using allozymes. The results showed that four (30.7%) of the 13 loci were polymorphic within the species, while the mean percentage of polymorphic loci within the populations was 1.9%. The mean number of alleles per locus for the species was 1.8 and 1.02 across the populations. Genetic diversity at the species level was low (H _es=0.135), and mean population diversity was notably lower (H _ep=0.005). A high degree of genetic differentiation was observed among populations. The salient points emerging from this study are: (1) statistically significant differences were found in allele frequencies among populations for every polymorphic locus (P<0.001), (2) the high mean coefficient of gene differentiation (G _ST) showed that 95% of the total allozyme variation was attributable to differences among populations, and (3) relatively high genetic distances between the populations were obtained (mean D=0.16). The Norwegian populations had the highest genetic diversity as compared with the other populations. Geographical comparisons revealed three different groups of populations clearly differentiated, i.e. Scandinavia (Norway and Sweden), Iceland and Russia. Cluster and principal coordinates analyses revealed the same genetic patterns of relationships among populations. Generally, this study indicates that E. alaskanus contains low allozymic variation in its populations. The implications of these results for the conservation of the species are discussed. Received: 23 October 1998 / Accepted: 19 December 1998     

Rapid evolutionary responses in a translocated population of intertidal snail (<Emphasis Type="Italic">Bembicium vittatum</Emphasis>) utilise variation from different source populations

The artificial movement of individuals between populations (translocation) can be an effective way to increase genetic diversity within populations, but few studies have undertaken long term genetic monitoring to determine if variation introduced by translocation is maintained over many generations or whether it can be used to adapt to local conditions. Here, we report on the changes in morphological and molecular variation over a 12-year period in a population of an intertidal littorine snail (Bembicium vittatum) that was created by mixing individuals from three geographically disjunct populations. These source populations differ genetically in shell shape and in allele frequency at several allozyme loci. We found that the translocated population had higher allozyme diversity than any of the source populations and that this pattern was maintained over multiple generations. Variation in shell shape also increased, but this declined over time as shells became taller. Some allozyme loci also showed significant changes in frequency over time. These changes were not consistently towards the genetic makeup of a single source population, and in the case of shell shape, were towards a phenotype that was most suited to the local environment. Our results suggest that genetic variation introduced into a population by translocation can be rapidly incorporated and used to adapt to local conditions without domination by a single source population’s genome. However, more studies are needed before generalisations on the benefits of mixing individuals from disjunct populations can be made.     

Spatial and demographic population genetic structure in Catasetum viridiflavum across a human-disturbed habitat

Spatial and temporal genetic structures were examined across sites on islands and mainland (continuous forest) populations of an epiphytic orchid, Catasetum viridiflavum, using 17 polymorphic allozyme loci. I tested whether patches on islands or at mainland sites comprised small local populations or a large population. Low among population differentiation was observed across the landscape suggesting that the species-specific pollinator and tiny wind-dispersed seeds maintain interconnections among distant patches. Temporal genetic structure among stage classes, and among breeding individuals are important components of the maintenance of genetic variation in this orchid. The natural history of this species including small breeding populations, probable high frequency of mating among relatives, and the high rates of seed movement among sites contribute to the high FIS. These data show that physically isolated patches in this epiphytic orchid comprise a single larger genetic population, which is independent of the physical distances among sites. Although quite different in ecological and life history characteristics, the genetic structure of this orchid demonstrates a pattern similar to temperate and tropical trees in fragmented landscapes.     

Microsatellite variation in natural Drosophila melanogaster populations from New South Wales (Australia) and Tasmania

Microsatellite variation was studied at 48 microsatellite loci in 10 Drosophila melanogaster populations to investigate the population structure on the Australian east coast. Low, but statistically significant population differentiation was observed among most populations. The populations on the Australian mainland did not show evidence for isolation by distance. We conclude that the population structure of D. melanogaster on the Australian mainland is probably the result of a shared history (recent colonization). The observed differences between local D. melanogaster populations probably reflect variation in effective population sizes rather than patterns of gene flow. Two populations from Tasmania were more differentiated from the Australian mainland than a population from Israel, raising the question whether they are derived from the Australian mainland or colonized from a different source population.     

Genetic variation at enzyme loci in North Atlantic minke whales,Balaenoptera acutorostrata

Electrophoretic variation within and between North Atlantic minke whale samples(Balaenoptera acutorostrata) from West Greenland, Iceland, and Norway was investigated. In the West Greenland samples, 28 enzyme systems were examined, representing 36 loci, of which 6 were found to be polymorphic. In Icelandic and Norwegian samples, 22 enzyme systems were examined, representing 29 loci, of which 6 and 5 were found to be polymorphic, respectively. The average heterozygosity was 0.058 (SE=0.024) in samples from West Greenland, 0.074 (SE=0.028) in samples from Iceland, and 0.054 (SE=0.023) in samples from Norway. No significant deviations from the expected Hardy—Weinberg genotypic frequencies, within samples taken from the same area, were found. Significant differences in allele frequencies were observed, however, between samples from the three different areas. The average Nei's genetic distance was 0.014 and the averageF _st value was 0.126. The genetic differences between the samples from the different areas indicate that those from West Greenland, Iceland, and Norway represented different breeding populations.     

黑头酸臭蚁微卫星标记的分离与开发(英文)

【目的】本研究旨在分离黑头酸臭蚁Tapinoma melanocephalum基因组微卫星标记,确定这些微卫星位点的多态性。【方法】使用454 GS-FLX焦磷酸测序技术开发来自中国华南陆地和岛屿的11个黑头酸臭蚁地理种群基因组微卫星位点。从随机设计的100对微卫星引物中筛选出10对引物,用于确定黑头酸臭蚁4个地理种群［东澳岛(DAD)、荷包岛(HBD)、梅州(MZ)和山咀(SJ)］10个微卫星位点的多态性,分析种群遗传多样性和种群分化。【结果】从11个黑头酸臭蚁地理种群基因组中成功开发和分离10对微卫星引物。在DAD, HBD, MZ和SJ 4个地理种群中,10个微卫星位点中7个有高多态性,这10个位点均显著偏离Hardy-Weinberg平衡;每个位点的等位基因数量(A)是3.50~9.00个,每个地理种群每个位点等位基因丰富度(A_R)在1.992~12.938之间。岛屿地理种群(DAD和HBD)的AR和预期杂合度(H_E)与大陆地理种群(MZ和SJ)的相比差异不显著。4个地理种群均显示高水平遗传分化(F_ST=0.15969);HBD和MZ种群与其他配对地理种群相比,遗传分化较高(F_ST=0.185),基因流较低,说明这两个种群基因流被限制。此外,遗传变异来自种群内个体之间。【结论】筛选新的微卫星位点能够为研究黑头酸臭蚁种群结构和繁殖结构提供有效工具,以深入了解其传播机制。     

Comparison of population genetic diversity between a rare, narrowly distributed species and a common, widespread species of Alnus (Betulaceae)

Comparisons of population genetic diversity between related rare and widespread species provide valuable insights to the consequences of rarity and are critical for conservation planning. Population genetic diversity of A. maritima, a rare species, was compared with its common, widespread congener A. serrulata to evaluate the impacts of small population size and high isolation on genetic diversity in A. maritima and to provide population genetic data to be used in conservation planning for A. maritima. Genetic data were also used to evaluate whether the disjunct distribution of A. maritima was due to range reduction or anthropogenic dispersal. Genetic diversity was lower in A. maritima (H(e) = 0.217) than in A. serrulata (H(e) = 0.268), and there is also higher inbreeding within A. maritima populations (f = 0.483) than A. serrulata populations (f = 0.269). The partitioning of genetic variation was also higher among A. maritima populations (Θ = 0.278), but not significantly different from that of A. serrulata (Θ = 0.197). Significant genetic differences among A. maritima populations support using local populations as seed sources for regional conservation efforts. The results also indicate that the highly disjunct distribution of A. maritima is due to natural range reduction in the past and not anthropogenic establishment of Oklahoma and Georgia populations.     

Large mainland populations of South Island robins retain greater genetic diversity than offshore island refuges

For conservation purposes islands are considered safe refuges for many species, particularly in regions where introduced predators form a major threat to the native fauna, but island populations are also known to possess low levels of genetic diversity. The New Zealand archipelago provides an ideal system to compare genetic diversity of large mainland populations where introduced predators are common, to that of smaller offshore islands, which serve as predator-free refuges. We assessed microsatellite variation in South Island robins (Petroica australis australis), and compared large mainland, small mainland, natural island and translocated island populations. Large mainland populations exhibited more polymorphic loci and higher number of alleles than small mainland and natural island populations. Genetic variation did not differ between natural and translocated island populations, even though one of the translocated populations was established with five individuals. Hatching failure was recorded in a subset of the populations and found to be significantly higher in translocated populations than in a large mainland population. Significant population differentiation was largely based on heterogeneity in allele frequencies (including fixation of alleles), as few unique alleles were observed. This study shows that large mainland populations retain higher levels of genetic diversity than natural and translocated island populations. It highlights the importance of protecting these mainland populations and using them as a source for new translocations. In the future, these populations may become extremely valuable for species conservation if existing island populations become adversely affected by low levels of genetic variation and do not persist.