Phenotypic plasticity inCrepis tectorum (Asteraceae)

Two experiments were carried out using two different approaches to compare populations ofCrepis tectorum (Asteraceae). One was based on a comparison of means of various vegetative and reproductive characters and another was based on a comparison of response patterns of the same characters in a series of environments. Population divergence within two earlier recognized form series, one from weed habitats and one from alvar habitats on Baltic islands, resulted in a partially overlapping pattern in cluster analyses based on character means. However, the pattern revealed by a comparison of the direction and amount of plastic response suggested that populations within the two form series had more similar response patterns than other combinations of populations. It was concluded that patterns of plasticity may provide useful additional information on the overall similarity among taxa. An hypothesis that plants in weed populations should exhibit a greater phenotypic response to the environments than plants in alvar populations was rejected.     

The evolution of self-fertility inCrepis tectorum (Asteraceae)

The extent of self-fertility was examined in 16 populations ofCrepis tectorum. A hypothesis that a weedy habit favours autogamy was only partly supported. Low levels of self-fertility characterized non-weedy populations from calcareous grasslands (alvars) on the Baltic island in Öland. By contrast, plants in nearly all weed populations studied were more or less self-fertile. However, the trend towards autogamy may have occurred independently of the trend towards a weedy habit, as shown by moderately to high levels of self-fertility in alvar populations from two other Baltic islands. In the weed group, there was a tendency for plants from two field populations to be more autogamous than plants from more ruderal habitats. There was an association between self-fertility and small, inconspicious heads in the alvar group but the association was weaker when weed populations were also considered. The relatively wide heads characterizing the ruderal weed populations may, at least partly, be an indirect effect of increases in overall plant size and/or in the size of the fruit associated with each flower.     

Genetic structure in the species-pair Silene vulgaris and S. uniflora (Caryophyllaceae) on the Baltic island of Öland

Allozyme data were used to assess the genetic structure between 37 sympatric populations of the species-pair Stlene vulgaris and S uniflora ssp petraea, and to infer levels of intra- and interspecific gene flow in the two species Silene vulgaris is a geographically widespread weed of disturbed habitats whereas S uniflora ssp petraea is endemic to the Baltic islands of Oland and Gotland On Oland, Silene vulgaris forms extensive linear populations along roads while S uniflora ssp petraea occurs in sparse and spatially-separated populations in open limestone habitats Despite the differences in population size and structure between the two species, both species show extremely low levels of between-population differentiation Between-site differences account for <2% of the total allozyme diversity within Oland in S vulgaris, and < 1% in S uniflora ssp petraea Indirect estimates of gene flow are high for both species (Nm = 11 and 27, respectively) There is no relationship between genetic distance and geographic distance within either species, and the lack of genetic structure is consistent with the pollination biology of the species - both of which are predominantly moth-pollinated The two species hybridize in intermediate habitats, and the geographic distribution of species-characteristic alleles indicates a potential for spatially extensive interspecific gene flow Nevertheless, there are significant differences in allele frequencies between the two species and multivariate analyses show no overlap between populations of the two species The species are ecologically separated by their different habitat preferences and by differences in their flowering phenology There is no evidence that the endemic S uniflora ssp petraea is threatened by genetic contamination or assimilation by the widespread weed, S vulgaris     

Geographical variation and genetic analysis of leaf shape inCrepis tectorum (Asteraceae)

Canonical variate analysis of plants raised in a uniform environment was used to study the pattern of geographical variation in leaf shape ofCrepis tectorum (Asteraceae). The diversity in leaf shape was much greater among populations confined to areas with exposed bedrock in the Baltic region than among weed populations scattered throughout Europe and Canada. A Ward's clustering linked outcrop populations from the archipelago of SW. Finland and the islands of Öland (Sweden) and Saaremaa (Estonia) due to the deeply lobed leaves characterizing these populations, while outcrop populations along the coast of E. Sweden were grouped due to their weakly lobed, narrow and dentate leaves. Most of the weed populations were grouped together but there was no tendency for the variation in this group to be related to habitat or geographical location. A mosaic of variation reflected in sharp (random) differentiation among local populations was superimposed on the large-scale ecogeographical pattern.—Crossing data indicated that most of the variation in degree of leaf dissection is governed by one major gene with deeply lobed leaves dominant over weakly lobed leaves. I suggest that the simple pattern of inheritance may have favoured rapid evolutionary changes in leaf shape, particularly in the Baltic area which emerged relatively late from the sea. Genetic correlations may have constrained the pattern of variation at higher taxonomic levels, since some of the trait associations detected in a segregating F₂ generation were also found at the among-population level.     

Threatened Alvar Grasslands in NW Russia and their Relationship to Alvars in Estonia

Alvar grasslands are thin-soil calcareous grasslands on Silurian and Ordovician limestone bedrock. They are considered unique plant communities for northern Europe and America with a high conservation value. Until now, European alvar grasslands have only been described in Sweden and Estonia. We described species-rich alvar grasslands on the Izhora plateau, NW Russia and compared this vegetation with that of Estonian alvar grasslands. Floristic composition and environmental conditions of Russian alvars were similar to Estonian alvar grasslands on thicker dry soil. Russian alvars are isolated from similar communities in Estonia, but extensive transport of domestic animals from Estonia to NW Russia in past centuries has probably enhanced species dispersal. Some characteristic Estonian alvar species were missing in NW Russia because it lies outside their natural ranges. Other differences in vegetation were explained by higher atmospheric nitrogen deposition and younger age of Russian sites. In NW Russia the alvar grasslands were small, but several red-listed plant species were present. In order to preserve European alvar vegetation effectively, Russian alvar grasslands need further study and conservation as an outpost of this threatened vegetation type.     

Large plant size counteracts early seed predation during the extended flowering season of a Silene uniflora (Caryophyllaceae) population

A population of the alvar race of the perennial herb, Silene uniflora (Caryophyllaceae), which grows on thin soils in open alvar habitats on the Baltic island of Oland SE Sweden, was found to have an extended and more or less bimodal flowering phenology Large individuals produced flowers during both periods, while small, and presumably young, individuals only produced flowers in either of the two periods, or in part of both In the early flowering period plants were heavily infested by the seed-predatory larvae of a noctuid moth, Hadena confusa , but in the late flowering period only a small proportion of the fruits was attacked by the seed predators The proportion of flowers developing into fruits was consistently high throughout the season Both the number of seeds per capsule and the mass of seeds decreased over the flowering season However, the germination success of early and late seeds did not differ Thus, although differing in number of seeds, both early and late flowers contributed to individual reproductive success Large individuals started to flower early in the season and despite their high loss of seeds in the early part of the summer they contributed a larger number of seeds to the seed pool than smaller and later-flowering individuals Although selection was acting to favour later flowering during a year with high early seed predation, consistency of date of flower initiation and of relative predation impact on individuals across years was low suggesting that recurrent selection by seed predators is weak Seed predation, although heavy, is therefore judged to be unlikely to cause a significant evolutionary response on flowering phenology in this plant     

Cranial variation in the European badger Meles meles (Carnivora,Mustelidae) in Scandinavia

Morphometric variation in 30 craniometric characters of 465 skulls of the European badgers (Meles meles) from across Europe was analysed. Multivariate analyses revealed that the populations from Norway, Sweden, and Finland differ from other European populations in having smaller skulls. The analyses also revealed significant differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ forms. On average, badgers from south‐west Norway were smaller than those of the remaining Fennoscandia. Morphological differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ populations of M. meles suggest a possible in situ semisympatric divergence since the beginning of the Holocene warming, or a complex history of two groups involving at least two colonization routes. The small‐sized Scandinavian badgers may be close to the ancestral form that used to be widespread in Denmark and throughout Europe. The animals from south‐west Norway may instead be descendants of ancestors that were the first to penetrate the southern parts of the Scandinavian Peninsula. The ‘main Fennoscandian’ badgers are likely to have been the descendants of the second wave of recolonization of Scandinavia. Specifically, they might have colonized the Scandinavian Peninsula from the east after the last glaciation.     

Population genetic consequences of geographic disjunction: a prairie plant isolated on Great Lakes alvars

Species may often exhibit geographic variation in population genetic structure due to contemporary and historical variation in population size and gene flow. Here, we test the predictions that populations on the margins of a species' distribution contain less genetic variation and are more differentiated than populations towards the core of the range by comparing patterns of genetic variation at five microsatellite loci between disjunct and core populations of the perennial, allohexaploid herb Geum triflorum. We sampled nine populations isolated on alvar habitat within the eastern Great Lakes region in North America, habitats that include disjunct populations of several plant species, and compared these to 16 populations sampled from prairie habitat throughout the core of the species' distribution in midwestern Canada and the USA. Alvar populations exhibited much lower within-population diversity and contained only a subset of alleles found in prairie populations. We detected isolation by distance across the species' range and within alvar and prairie regions separately. As predicted, genetic differentiation was higher among alvar populations than among prairie populations, even after controlling for the geographic distance between sampled populations. Low diversity and high differentiation can be accounted for by the greater contemporary spatial isolation of alvar populations. However, the genetic structure of alvar populations may also have been influenced by postglacial range expansion and contraction. Our results are consistent with alvar populations being founded during an expansion of prairie habitat during the warmer, hypsithermal period approximately 5000 bp and subsequently becoming stranded on isolated alvar habitat as the climate grew cooler and wetter.     

Specialist taxa restricted to threatened habitats contribute significantly to the regional diversity of Peltigera (Lecanoromycetes,Ascomycota) in Estonia

The widespread cyanolichen genus Peltigera comprises many insufficiently known poorly delimited and/or undescribed species. Phylogenetic analysis of 252 Peltigera specimens from a wide range of habitat types in Estonia revealed 31 putative taxa (OTUs). Multivariate analysis revealed habitat-specific segregation between the Peltigera species along a gradient from humid eutrophic forests to dry oligotrophic forests and grasslands and along a soil pH gradient from alkaline soils of alvar grasslands to acidic soils of conifer forests. The diversity of Peltigera was the highest on roadsides and dunes and the lowest in alvar habitats which, however, supported the unique assemblage of undescribed Peltigera taxa. Deciduous broad-leaved forests, too, included several undescribed or rare and red-listed species. The results demonstrate that in Estonia many Peltigera species have narrow habitat requirements and are at present threatened by habitat loss and degradation.     

Morphometric differentiation,patterns of interfertility,and the genetic basis of character evolution inCrepis tectorum (Asteraceae)

Morphometric analyses of populations ofCrepis tectorum raised under uniform conditions support the earlier finding based on herbarium studies that extensive divergence has taken place in western Europe with a centre of diversity in the Baltic lowland area. A crossing experiment showed that subsp.pumila on the island of Öland has acquired a wide range of derived characters without the establishment of strong reproductive barriers, the only exception being a weak tendency for crosses with other populations on Öland to yield more fertile F₁ offspring than crosses with weedy and non-weedy populations in adjacent regions. Canadian weed populations showed a more uniform response in crosses with subsp.pumila than did morphologically similar weed populations scattered throughout Central Europe. While F₂ data suggest a monogenic basis of differences in the extent of leaf dissection, other distinctive traits appear to be governed by multiple genetic factors with individually slight effects.     

Genetic differentiation of southeast Baltic populations of sea trout inferred from single nucleotide polymorphisms

Sea trout (Salmo trutta m. trutta) is a migratory form of brown trout common in the Baltic Sea. Nine populations from the southeast Baltic (Poland; Lithuania; Denmark, Bornholm; Estonia and Russia) were genotyped using iPLEX Gold technology (Sequenom) with 62 informative SNPs. A diagnostic panel of 23 SNPs was applied to estimate genetic differentiation and assess the population structure of Baltic sea trout. The highest level of pairwise F_ST differences was observed between the Russian (East Gulf of Finland) and Polish (Baltic main basin) populations. The lowest differences were between the two Polish and the Polish and Lithuanian populations. A genetic similarity was noted between the Estonian Riguldi River and Danish Bornholm populations, and this finding was supported by a Bayesian and factorial correspondence analysis. Diversity within populations was highest for populations from Estonia and lowest for the Lithuanian population. Genetic structure analysis indicated that individuals from the nine populations were clustered into four groups.     

Island biogeography of North European parthenogenetic Lumbricidae: I. Clone pool affinities and morphometric differentiation of Åland populations

Genetic and morphometnc differences between island (Åland) and mainland (Sweden, Finland and Estonia) populations of five parthenogenetic and ecologically different Lumbricidae species were studied by means of enzyme electrophoresis and morphometric analysis Affinities of clones to the mainland populations show that the postglacial rates of clone flow to Åland differ widely among species In Octolasion cyaneum, the only clone found on Åland is not related to any of the studied clones from the nearby mainlands The Åland population evidently originates from one long dispersal jump through human agency The Åland clone pool of the anthropochorous Aporrectodea rosea is rich As many as 98% of the worms and 82% of the clones belong to genotypes found in the mainland populations Åland is an intergradation zone of A rosea clones that have arrived from the west (Sweden) and from the east (Finland and Estonia) The more intense cultural connections to Sweden explain the higher clonal affinities that A rosea in Åland shows to the west than to the east There are not many Octolasion tyrtaeum clones on Åland The few that are present are mainly clones also found on the mainlands On Åland Eisemella tetraedra has extensive diversity but low clonal affinities to the mainland populations This also holds true for Dendrobaena octaedra but clonal affinities are on an average higher than in E tetraedra The Aland clone pool of D octaedra shows more affinities to Finland than to the other mainlands Probably the main factor that has facilitated D octaedra clone flow from Finland is a continuous archipelago In most species there are at least some clones that are more abundant on Åland than on the mainlands They may have a wider niche in the island environment The Åland populations of A rosea, O tyrtaeum and D octaedra differed particularly in somatic traits from the mainland populations The secondary reproductive traits of the parthenogens on Åland did not differ from the mainland populations We conclude that except for O cyaneumÅland has a role as a stepping stone island in the postglacial dispersal of parthenogenetic earthworms across the Baltic Sea Different rates of flow by immigrant clones from the surrounding mainland populations have resulted in very dissimilar clone pool structures even on the same island The Åland populations also show some morphometric differences to the mainland populations     

The impact of exotic weed competition on a rare New Zealand outcrop herb,

Nearly one quarter of New Zealand’s unique vascular plant flora is threatened, and weed invasion is implicated in the decline of more than half of these threatened species. However, there is little experimental evidence showing that invasive weeds have a direct impact on threatened native plants. This study experimentally tested the hypothesis that competition with invasive weeds threatens the rare outcrop plant Pachycladon cheesemanii (Brassicaceae). Pachycladon cheesemanii is a threatened South Island, New Zealand endemic with a distribution nearly confined to rock outcrops. It has disappeared from historical record sites throughout its range. The effects of weed competition and habitat on P. cheesemanii establishment, growth and survival were investigated by sowing seed into replicated plots subject to three treatments: weed removal, soil disturbed and unweeded control, in three habitat types: forested and open rock outcrops and open tussock grassland. The experiments were carried out at three locations: Mt Somers (Canterbury), Wye Creek and Diamond Lake (Otago). Within weedy rock outcrop habitat, weed removal significantly increased the rate of P. cheesemanii germination, and appeared to increase seedling growth rates, implying that weeds can negatively impact populations. Relative to rock outcrop habitat, P. cheesemanii germination was very low in adjacent open grassland habitat regardless of weeding treatment. Demographic monitoring of four natural populations of P. cheesemanii revealed that seed production is highly variable among populations and may be limited by browse and mechanical damage to inflorescences. Pachycladon cheesemanii does produce a persistent seed bank but most seed is found close to parent populations. Our results suggest that competition with invading weeds threatens current P. cheesemanii populations, that plant establishment can be enhanced by weed removal, and that considerable potential exists for artificially expanding populations by sowing seed into appropriate weed-free habitat.     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

Native perennial grasses show evolutionary response to Bromus tectorum (cheatgrass) invasion

Invasive species can change selective pressures on native plants by altering biotic and abiotic conditions in invaded habitats. Although invasions can lead to native species extirpation, they may also induce rapid evolutionary changes in remnant native plants. We investigated whether adult plants of five native perennial grasses exhibited trait shifts consistent with evolution in response to invasion by the introduced annual grass Bromus tectorum L. (cheatgrass), and asked how much variation there was among species and populations in the ability to grow successfully with the invader. Three hundred and twenty adult plants were collected from invaded and uninvaded communities from four locations near Reno, Nevada, USA. Each plant was divided in two and transplanted into the greenhouse. One clone was grown with B. tectorum while the other was grown alone, and we measured tolerance (ability to maintain size) and the ability to reduce size of B. tectorum for each plant. Plants from invaded populations consistently had earlier phenology than those from uninvaded populations, and in two out of four sites, invaded populations were more tolerant of B. tectorum competition than uninvaded populations. Poa secunda and one population of E. multisetus had the strongest suppressive effect on B. tectorum, and these two species were the only ones that flowered in competition with B. tectorum. Our study indicates that response to B. tectorum is a function of both location and species identity, with some, but not all, populations of native grasses showing trait shifts consistent with evolution in response to B. tectorum invasion within the Great Basin.     

Diversity of Aphanizomenon flos-aquae (cyanobacterium) populations along a Baltic Sea salinity gradient

Colony-forming cyanobacteria of the genus Aphanizomenon form massive blooms in the brackish water of the Baltic Sea during the warmest summer months. There have been recent suggestions claiming that the Baltic Sea Aphanizomenon species may be different from Aphanizomenon flos-aquae found in lakes. In this study, we examined variability in the morphology and 16S-23S rRNA internal transcribed spacer (ITS) sequences of A. flos-aquae populations along a salinity gradient from a string of lakes to a fjord-like extension of the Baltic Sea to the open Baltic Sea. Morphological differences among the populations were negligible. We found that the Baltic Sea was dominated (25 out of 27 sequences) by one ITS1-S (shorter band of ITS 1 [ITS1]) genotype, which also was found in the lakes. The lake populations of A. flos-aquae tended to be genetically more diverse than the Baltic Sea populations. Since the lake ITS1-S genotypes of A. flos-aquae are continuously introduced to the Baltic Sea via inflowing waters, it seems that only one ITS1 genotype is able to persist in the Baltic Sea populations. The results suggest that one of the ITS1-S genotypes found in the lakes is better adapted to the conditions of the Baltic Sea and that natural selection removes most of the lake genotypes from the Baltic Sea A. flos-aquae populations.     

Restoration of alvar vegetation on Öland,Sweden

Abstract. Alvars in the Baltic region, particularly on the Swedish islands of Öland and Gotland and in western Estonia, are well‐known for their plant species richness and extensive populations of rare species. Grasslands make up most of alvar vegetation. The extent of these grasslands decreases because of bush enchroachment which takes place in most alvars when traditional land use practice changes, notably when grazing is ceased. The main threat for the alvar grassland is formed by the rapid expansion of Juniperus communis and Potentilla fruticosa in dry and wet sites, respectively. Applied research has been carried out during several years in order to develop plans for the restoration of alvar grassland. Thus it was shown that species richness in areas with a mixture of grassland and shrubland decreases in relation to the increased cover by shrubs. In the case of Juniperus there is a distinct drop in species number as soon as the cover exceeds 75‐80%, while the decrease is more gradual with increased cover of Potentilla. The seed banks under dense stands of these shrubs only contain ca. 20% of the species found in dry and wet alvar meadows. This was confirmed by clearing experiments. Long‐term recordings have shown that establishment of juniper seedlings takes place mainly in half‐open areas between already existing junipers. Intermediate‐sized junipers expanded faster than old and big shrubs. Potentilla shrubs recover fast from cutting or burning. After 2‐4 yr they have almost attained their former size. This recovery can be prevented when cattle grazing is introduced. In the framework of a local alvar restoration project on Öland, covering 7000 ha, grazing regimes are re‐established, fences erected and large‐scale clearings carried out. Recently established juniper shrubs are being cleared, intermediate sized junipers (30‐50 yr old) are selectively removed while creating a mosaic landscape with high biological diversity. Older dense juniper stands are left alone or are only partly cleared. Potentilla stands in moist areas are cut to create moist meadows which are breeding grounds for waders, and to establish corridors between remaining open areas. Three items are discussed (1) the importance of the Stora Alvar area; (2) re‐introduction of grazing and (3) re‐introduction of species. The outstanding importance of the area regarding species richness and endemism should be recognized through a ‘golden list’ to be used along with red and blue lists.     

Variation in heteroblastic succession among populations of Crepis tectorum

Eleven populations of the monocarpic species, Crepis tectorurn (Asteraceae) in South Sweden differed in the extent to which different leaf characters changed during plant ontogeny. Multivariate analyses on sequences of leaf samples collected from greenhouse-grown plants of different ages revealed a group structure of populations that was different from that revealed by analysing variation in leaves from adult plants. Ontogenetic data suggested that populations within a weed and an alvar ecotype constituted more natural groups than combinations of populations from both eco-types that had similar leaves at adult stages, supporting the hypothesis that these ecotypes may have different migration histories in this area. Variation among the populations in leaf characteristics was partly due to differences that were present at all ontogenetic stages and partly due to variation in the rate of development of some of the leaf characters during plant ontogeny. Plants in some populations appeared to have more 'juvenile' leaves at adult stages than plants in other populations, at least in some characters.     

Conservation genetics and phylogeography of southern dunlins Calidris alpina schinzii

Breeding populations of southern dunlin Calidris alpina schinzii in South Fennoscandia and the Baltic are severely fragmented and declining dramatically. Information on the genetic structure and diversity is therefore of importance for the conservation and management of these populations. Here we present the results of comparative genetic analyses of these populations with other populations of the schinzii , alpina and arctica subspecies in northern Europe. We sequenced the mitochondrial DNA control region and the Z-chromosome intron VLDLR-9, and analyzed microsatellites and AFLPs, for analyses of within-population genetic diversity. We also extended previous analyses of the phylogeographic structure of dunlins in northern Europe with a larger sample of individuals and populations. Our results revealed no evidence of reduced genetic diversity or increased levels of inbreeding in the small and fragmented populations around the Baltic Sea as compared to the more vital and larger populations elsewhere. Nevertheless, their small population sizes and presumably high degree of isolation may lead to local extinctions, indicating that demographic and ecological factors may pose a greater threat to the survival of these populations than purely genetic factors. Phylogeographically, the schinzii populations in Scandinavia and the Baltic do not form a separate genetic clade, but are part of larger cline of genetic variation encompassing several recognized subspecies of dunlins in the western Palearctic region. Only the Icelandic population showed some distinctiveness in genetic structure and might therefore be considered a separate management unit. Our study highlights the general problem of lack of genetic support for subspecies in avian taxonomy and conservation genetics.