Genetic admixture is supposed to be an important trigger of species expansions because it can create the potential for selection of genotypes suitable for new climatic conditions. Up until now, however, no continent-wide population genetic study has performed a detailed reconstruction of admixture events during natural species expansions. To fill this gap, we analysed the postglacial history of Alnus glutinosa, a keystone species of European swamp habitats, across its entire distribution range using two molecular markers, cpDNA and nuclear microsatellites. CpDNA revealed multiple southern refugia located in the Iberian, Apennine, Balkan and Anatolian Peninsulas, Corsica and North Africa. Analysis of microsatellites variation revealed three main directions of postglacial expansion: (i) from the northern part of the Iberian Peninsula to Western and Central Europe and subsequently to the British Isles, (ii) from the Apennine Peninsula to the Alps and (iii) from the eastern part of the Balkan Peninsula to the Carpathians followed by expansion towards the Northern European plains. This challenges the classical paradigm that most European populations originated from refugial areas in the Carpathians. It has been shown that colonizing lineages have met several times and formed secondary contact zones with unexpectedly high population genetic diversity in Central Europe and Scandinavia. On the contrary, limited genetic admixture in southern refugial areas of A. glutinosa renders rear-edge populations in the Mediterranean region more vulnerable to extinction due to climate change.

Background/AimsRecently, new palaeoecological records supported by molecular analyses and palaeodistributional modelling have provided more comprehensive insights into plant behaviour during the last Quaternary cycle. We reviewed the migration history of species of subgenus Alnus during the last 50,000 years in Europe with a focus on (1) a general revision of Alnus history since the Last Glacial Maximum (LGM), (2) evidence of northern refugia of Alnus populations during the LGM and (3) the specific history of Alnus in particular European regions.MethodologyWe determined changes in Alnus distribution on the basis of 811 and 68 radiocarbon-dated pollen and macrofossil sites, respectively. We compiled data from the European Pollen Database, the Czech Quaternary Palynological Database, the Eurasian Macrofossil Database and additional literature. Pollen percentage thresholds indicating expansions or retreats were used to describe patterns of past Alnus occurrence.Principal FindingsAn expansion of Alnus during the Late Glacial and early Holocene periods supports the presence of alders during the LGM in southern peninsulas and northerly areas in western Europe, the foothills of the Alps, the Carpathians and northeastern Europe. After glaciers withdrew, the ice-free area of Europe was likely colonized from several regional refugia; the deglaciated area of Scandinavia was likely colonized from a single refugium in northeastern Europe. In the more northerly parts of Europe, we found a scale-dependent pattern of Alnus expansion characterised by a synchronous increase of Alnus within individual regions, though with regional differences in the times of the expansion. In southern peninsulas, the Alps and the Carpathians, by contrast, it seems that Alnus expanded differently at individual sites rather than synchronously in whole regions.ConclusionsOur synthesis supports the idea that northern LGM populations were important sources of postglacial Alnus expansion. The delayed Alnus expansion apparent in some regions was likely a result of environmental limitations.

Newly discovered tetraploid populations are situated in the putative main glacial refugia, and neither of them was likely to have been involved in the colonization of central and northern Europe after glacial withdrawal. This could mean that neither the Iberian Peninsula nor the western part of the Balkan Peninsula served as effective refugial areas for northward post-glacial expansion of A. glutinosa.

SummaryWe reconstructed the historical pattern of postglacial biogeographic range expansion of the boreal tree species Alnus incana in Europe.To assess population genetic structure and diversity, we performed a combined analysis of nuclear microsatellite loci and chloroplast DNA sequences (65 populations, 1004 individuals).Analysis of haplotype and microsatellite diversity revealed that southeastern refugial populations situated in the Carpathians and the Balkan Peninsula did not spread north and cannot be considered as important source populations for postglacial recolonization of Europe; populations in Eastern Europe did not establish Fennoscandian populations; populations in Fennoscandia and Eastern Europe have no unique genetic cluster, but represent a mix with a predominant cluster typical for Central Europe; and that colonization of Fennoscandia and Eastern Europe took place from Central Europe.Our findings highlight the importance of an effective refugium in Central Europe located outside classical southern refugia confirming the existence of northern refugia for boreal trees in Europe. The postglacial range expansion of A. incana did not follow the model established for Picea abies. Fennoscandian populations are not derived from Eastern European ones, but from Central European ones.

We attempted to confirm that seed banks can be viewed as an important genetic reservoir by testing the hypothesis that standing (aboveground) plants represent a nonrandom sample of the seed bank. We sampled multilocus allozyme genotypes from three species with different life history strategies: Amaranthus retroflexus, Carduus acanthoides, Pastinaca sativa. In four populations of each species we analysed the extent to which allele and genotype frequencies vary in consecutive life history stages including the summer seed bank, which has been overlooked up to now. We compared the winter seed bank (i.e., seeds collected before the spring germination peak), seedlings, rosettes, the summer seed bank (i.e., seeds collected after the spring germination peak) and fruiting plants. We found that: (1) All three species partitioned most of their genetic diversity within life history stages and less among stages within populations and among populations. (2) All genetic diversity parameters, except for allele frequencies, were similar among all life history stages across all populations in different species. (3) There were differences in allele frequencies among life history stages at all localities in Amaranthus retroflexus and at three localities in both Carduus acanthoides and Pastinaca sativa. (4) Allele frequencies did not differ between the winter and summer seed bank in most Carduus acanthoides and Pastinaca sativa populations, but there was a marked difference in Amaranthus retroflexus. In conclusion, we have shown that the summer seed bank is not genetically depleted by spring germination and that a majority of genetic diversity remains in the soil through summer. We suggest that seed banks in the species investigated play an important role by maintaining genetic diversity sufficient for recovery rather than by accumulating new genetic diversity at each locality.

Citation: Moravcová L, Pyšek P, Jarošík V, Zákravský P (2011) Potential phytotoxic and shading effects of invasive Fallopia (Polygonaceae) taxa on the germination of native dominant species. NeoBiota 9: 31-47. doi: 10.3897/neobiota.9.1266 Abstract Two species of the genus Fallopia (F. sachalinensis, F. japonica, Polygonaceae) native to Asia, and their hy brid (F. ×bohemica), belong to the most noxious plant invaders in Europe. They impact highly on invaded plant communities, resulting in extremely poor native species richness. The low number of native species in invaded communities points to the possible existence of mechanisms suppressing their germination. In this study we assessed, under laboratory conditions, whether there are phytotoxic effects of the three Fallopia congeners on seed germination of three target species: two native species commonly growing in habi tats that are often invaded by Fallopia taxa (Urtica dioica, Calamagrostis epigejos), and Lepidium sativum, a species commonly used in allelopathic bioassays as a control. Since Fallopia taxa form dense stands with high cover, we included varying light conditions as an additional factor, to simulate the effects of shading by leaf canopy on germination. The effects of aqueous extracts (2.5%, 5.0%, and 0% as a control) from dry leaves and rhizomes of the Fallopia congeners on germination of the target species were thus studied under two light regimes, simulating full daylight (white light) and light filtered through canopy (green light), and in dark as a control regime. Rhizome extracts did not affect germination. Light treatments yielded inconclusive results, indicating that poor germination and establishment of species in invaded stands is unlikely to be caused by shading alone. However, we found a pronounced phytotoxic effect of leaf extracts of Fallopia taxa, more so at 5.0% than 2.5% extract concentration. NeoBiotaLenka Moravcová et al. / NeoBiota 9: 31-47 (2011) 32 exerted the largest negative effect on the germination of Urtica dioica, F. ×bohemica on that of C. epigejos, and F. japonica had invariably the lowest inhibitory effect on all test species. The weak phytotoxic effect of F. japonica corresponds to the results of previous studies that found this species to be generally a weaker competitor than its two congeners. Although these results do not necessarily provide direct evidence for allelopathic effects in the field, we demonstrate the potential phytotoxic effect of invasive Fallopia taxa on the germination of native species. This suggests that allelopathy may play a role in the impact of Fallopia invasion on species diversity of invaded communities.

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