Phylogeny and quaternary history of the European montane/alpine endemicSoldanella(Primulaceae) based on ITS and AFLP variation

Soldanella contains 16 species of herbaceous perennials that are endemic to the central and south European high mountains. The genus is ecogeographically subdivided into forest/montane and alpine species. Evolutionary relationships and large-scale biogeographic patterns were inferred from parsimony analyses of the internal transcribed spacer (ITS) regions of nuclear ribosomal DNA, and genetic distance analyses based on amplified fragment length polymorphism (AFLP) markers. The ITS region proved useful for examining subgeneric relationships and testing hypotheses on genus-wide divergence times, whereas the AFLP markers were suitable for studying relationships among closely related taxa and b…

The effect of Quaternary climatic changes on plant distribution and evolution

Abstract Climatic oscillations in the Quaternary have played a major role in changing the geographical distribution of plant species. Recent molecular work has provided new insights into the location of glacial refugia and routes of postglacial expansion. A comparative analysis shows that phylogeographic patterns in Europe appear to be less congruent than in North America. The change of geographical distribution has also provided opportunities for speciation through isolation, and hybridization after secondary contact. However, a complete understanding of the significance of the Quaternary climatic changes for plant speciation requires further research into the estimation of reliable and ac…

Evolution in the Arctic: a phylogeographic analysis of the circumarctic plant,Saxifraga oppositifolia(Purple saxifrage)

Summary • A survey of chloroplast DNA variation in the circumarctic-alpine plant, Saxifraga oppositifolia, has resolved two highly divergent cpDNA lineages with geographically widespread and mainly allopatric distributions that are largely concordant with those of two subspecies, that is, subspp. oppositifolia and glandulisepala. These subspecies differ for a single morphological trait and hence level of morphological divergence does not equate to molecular divergence within the species. • The two cpDNA lineages were estimated to have diverged from their most recent common ancestor 5.37–3.76 Ma, that is, during the early to mid-Pliocene. • A nested clade analysis was conducted in an attempt…

Patterns of genetic and phenotypic variation in Iris haynei and I. atrofusca (Iris sect. Oncocyclus = the royal irises) along an ecogeographical gradient in Israel and the West Bank.

Iris haynei and I. atrofusca are two closely related narrow endemics distributed vicariously along an ecogeographical north-south gradient in Israel and the West Bank. To obtain baseline information of the taxonomic status, conservation and population history of these taxa, we investigated patterns of phenotypic variation and the partitioning of genetic variation within and among populations using dominant random amplified polymorphic DNA (RAPD) markers. Multivariate (principal components analysis) and taxonomic distance analyses based on morphometric traits from eight populations revealed no unambiguous separation into two distinct groups. Results of genetic analyses for nine populations d…

Late Quaternary distributional stasis in the submediterranean mountain plant Anthyllis montana L. (Fabaceae) inferred from ITS sequences and amplified fragment length polymorphism markers.

Anthyllis montana is a submediterranean, herbaceous plant of the southern and central European mountains. The internal transcribed spacer (ITS) regions of nuclear ribosomal DNA were sequenced from multiple accessions of the species and several closely related taxa. In addition, amplified fragment length polymorphism (AFLP) was analysed from 71 individuals of A. montana collected in 20 localities, mainly in the Pyrenees, Alps, Italian Peninsula and Balkans. Our ITS phylogeny showed a sequential branching pattern in A. montana, implying a western Mediterranean origin followed by an eastward migration. ITS clock calibrations suggest that speciation of A. montana took place at the Pliocene-Plei…

The genetic ghost of an invasion past: colonization and extinction revealed by historical hybridization inSenecio

Hybridization is an important evolutionary factor in the diversification of many plant and animal species. Of particular interest is that historical hybridization resulting in the origin of new species or introgressants has occurred between species now geographically separated by great distances. Here, we report that Senecio massaicus, a tetraploid species native to Morocco and the Canary Islands, contains genetic material of two distinct, geographically separated lineages: a Mediterranean lineage and a mainly southern African lineage. A time-calibrated internal transcribed spacer phylogeny indicates that the hybridization event took place up to 6.18 Ma. Because the southern African lineage…

Plant speciation in continental island floras as exemplified byNigellain the Aegean Archipelago

Continental shelf island systems, created by rising sea levels, provide a premier setting for studying the effects of geographical isolation on non-adaptive radiation and allopatric speciation brought about by genetic drift. The Aegean Archipelago forms a highly fragmented complex of mostly continental shelf islands that have become disconnected from each other and the mainland in relatively recent geological times (ca<5.2 Ma). These ecologically fairly homogenous islands thus provide a suitable biogeographic context for assessing the relative influences of past range fragmentation, colonization, gene flow and drift on taxon diversification. Indeed, recent molecular biogeographic studies…

Past, present and future of mountain species of the French Massif Central - the case ofSoldanella alpinaL. subsp.alpina(Primulaceae) and a review of other plant and animal studies

Aim  Our goals were: (1) to investigate patterns of genetic variation in the French Massif Central (MC) of Soldanella alpina (Primulaceae), an alpine plant species that has only one known population in the region; (2) to analyse these patterns in order to deduce the Quaternary history of the population and to predict how current climatic warming may affect it; and (3) to review molecular analyses from the MC to evaluate the importance of the region for the conservation of genetic diversity. Location  Europe, with a special focus on the French Massif Central and adjacent regions. Methods  Amplified fragment length polymorphisms (AFLPs) were analysed for 192 individuals (nine populations) of …

Genecological and isozyme studies in Senecio vernalis Waldst. & Kit. and S. vulgaris L. var. vulgaris (Asteraceae) from Central Europe and Israel

Summary Genecological evidence and evidence obtained from an isozyme survey suggest a considerable amount of intraspecific differentiation in Senecio vernalis . Plants of this species from Israel were found to differ strikingly from those of C European (German) origin in most of the traits investigated, e.g., in the possession of pronounced seed dormancy, which seems to be governed by a single gene, a shorter life cycle, lack of rosette growth correlated with stem elongation prior to budding, a larger number of smaller capitula with fewer (disc) florets, and an ephemeral habit with plants only flowering soon after the first winter rains. Both accessions achieved a similar reproductive poten…

Late Quaternary history ofHippophaë rhamnoidesL. (Elaeagnaceae) inferred from chalcone synthase intron (Chsi) sequences and chloroplast DNA variation

Fossil pollen records indicate that Hippophae rhamnoides (Sea Buckthorn) was widespread on late- and early postglacial raw soils throughout much of central and northern Europe, but that Early Holocene reforestation restricted populations to northern coastal habitats, or along mountain streams in the Alps, Pyrenees, and Carpathians. We used sequence variation at the nuclear chalcone synthase intron ( Chsi ), in conjunction with chloroplast DNA–restriction fragment length polymorphism data, to investigate the intraspecific phylogeny, phylogeographic structure, and expansion demographic history of this dioecious and wind-pollinated shrub at its range-wide scale in Europe and Asia Minor. Four m…

Chloroplast DNA and isozyme analysis of the progenitor-derivative species relationship between Senecio nebrodensis and S. viscosus (Asteraceae )

A comparison of chloroplast DNA (cpDNA) and isozyme diversity in Senecio nebrodensis L., a species restricted to certain mountain ranges in Spain, and S. viscosus L., a widespread ruderal species in Europe, revealed that S. viscosus possessed the more common of two cpDNA haplotypes resolved in S. nebrodensis and contained only a small subset of the alleles found in S. nebrodensis at enzyme coding loci. The two species shared one restriction length mutation and one site mutation in their cpDNA, which distinguished them from other European Senecio species examined previously. Taken overall, these results support the hypothesis that S. nebrodensis and S. viscosus are related as a progenitor-de…

The Mediterranean region - a hotspot for plant biogeographic research.

Genetic basis of speed of development in Senecio vulgaris L var. vulgaris, S. vulgaris ssp. denticulatus (O.F. Muell.) P.D. Sell, and Senecio vernalis Waldst. & Kit.

The genetic basis of differences in speed of development from germination to first bud formation was investigated in Senecio vulgaris var. vulgaris and S. vulgaris ssp. denticulatus, and also in S. vernalis sampled from Israel and Germany. In the case of S. vulgaris, F2 segregation analysis and the recovery of very late and very early lines from extreme F2 phenotypes showed that differences can be explained by a single major gene model, whereas segregation data from F2 and backcross progenies in S. vernalis are not incompatible with a digenic model of inheritance. Senecio vernalis from Israel and S. vulgaris var. vulgaris reached the different developmental stages in a substantially shorter…

Quaternary diversification in European alpine plants: pattern and process

Molecular clock approaches applied previously to European alpine plants suggest that Primula sect. Auricula , Gentiana sect. Ciminalis and Soldanella diversified at the beginning of the Quaternary or well within this period, whereas Globularia had already started diversifying in the (Late–)Tertiary. In the first part of this paper we present evidence that, in contrast to Globularia and Soldanella , the branching patterns of the molecular internal transcribed spacer phylogenies of both Primula and Gentiana are incompatible with a constant–rates birth–death model. In both of these last two taxa, speciation probably decreased through Quaternary times, perhaps because of some niche–filling pro…

Causes of the genetic architecture of south-west European high mountain disjuncts

Background: Postglacial climatic warming in south-western Europe and the retreat of cold-adapted species into higher elevations, starting in the Sierra Nevada and proceeding northwards to the Pyrenees and Alps, should have resulted in a pattern of ‘successive vicariance’. Alternatively, long-distance dispersal might explain the extant distribution pattern of mountain species in this region. Aims: Here, we report an investigation of two alpine plants, Saxifraga oppositifolia and S. stellaris, which co-occur in the Sierra Nevada, the Pyrenees, and the south-western Alps/Massif Central. Our aim was to distinguish between (successive) vicariance and long-distance dispersal as alternative explan…

Chloroplast DNA and isozyme evidence on the evolution ofSenecio vulgaris (Asteraceae)

Chloroplast DNA (cpDNA) and isozyme variation were analyzed over a range of populations of two infraspecific taxa of the tetraploidSenecio vulgaris. The isozyme data were supportive of the hypothesis that the weedy and cosmopolitanS. vulgaris var.vulgaris is an evolutionary derivative ofS. vulgaris subsp.denticulatus from the coasts of W Europe and montane altitudes in S Spain and Sicily. The two taxa exhibited a very high genetic identity with subsp.denticulatus containing slightly more isozyme diversity than was found in var.vulgaris. — Three cpDNA haplotypes (A, B, C) already known from other Mediterranean diploid species ofSenecio were resolved in var.vulgaris, and an additional fourth …

Parallel bursts of recent and rapid radiation in the Mediterranean and Eritreo-Arabian biodiversity hotspots as revealed byGlobulariaandCampylanthus(Plantaginaceae)

An AFLP clock for the absolute dating of shallow-time evolutionary history based on the intraspecific divergence of southwestern European alpine plant species.

The dating of recent events in the history of organisms needs divergence rates based on molecular fingerprint markers. Here, we used amplified fragment length polymorphisms (AFLPs) of three distantly related alpine plant species co-occurring in the Spanish Sierra Nevada, the Pyrenees and the southwestern Alps/Massif Central to establish divergence rates. Within each of these species (Gentiana alpina, Kernera saxatilis and Silene rupestris), we found that the degree of AFLP divergence (D(N72)) between mountain phylogroups was significantly correlated with their time of divergence (as inferred from palaeoclimatic/palynological data), indicating constant AFLP divergence rates. As these rates d…

Reciprocal hybridization at different times between Senecio flavus and Senecio glaucus gave rise to two polyploid species in north Africa and south‐west Asia

Summary • The analysis of hybrid plant taxa using molecular methods has considerably extended understanding of possible pathways of hybrid evolution. • Here, we investigated the origin of the tetraploid Senecio mohavensis ssp. breviflorus and the hexaploid Senecio hoggariensis by sequencing of nuclear and chloroplast DNA, and by analysis of the distribution of taxon-specific amplified fragment length polymorphism (AFLP) fragments. • Both taxa originated from hybridization between the diploid Senecio flavus and Senecio glaucus. Whereas S. glaucus was the female parent in the origin of S. mohavensis ssp. breviflorus, S. flavus was the female parent in the origin of S. hoggariensis. • The dist…

Molecular inference of a Late Pleistocene diversification shift inNigellas. lat. (Ranunculaceae) resulting from increased speciation in the Aegean archipelago

Aim To infer the temporal course and geographical mode of speciation in Mediterranean/Southwest Asian Nigella s. lat. Location Mediterranean Basin, Aegean archipelago. Methods Phylogenies for Nigella L. and Garidella L. (= Nigella s. lat.) were obtained from maximum-likelihood analyses of internal transcribed spacer (ITS) sequences. Diversification through time was analysed by log lineages-through-time (LTT) plots and survival analyses. Relative node age estimates were regressed against the degree of sympatry between sister clades to infer the predominant mode of geographical speciation in Nigella s. lat. Results The Late Pleistocene radiation of the Nigella arvensis complex in the Aegean r…

Differential cycles of range contraction and expansion in European high mountain plants during the Late Quaternary: insights from Pritzelago alpina (L.) O. Kuntze (Brassicaceae).

Nuclear DNA sequence variation of the internal transcribed spacer (ITS) and amplified fragment length polymorphisms (AFLPs) were used to illuminate the evolutionary history of Pritzelago alpina, a herbaceous perennial of (sub)alpine to nival habitats of the European high mountains. Maximum likelihood analysis of ITS sequences of P. alpina, Hornungia petraea and Hymenolobus procumbens (the 'Pritzelago alliance') resolved P. alpina and H. petraea as sister taxa. ITS divergence estimates support an origin for P. alpina in the Late Tertiary, while intraspecific diversification started in the Late Quaternary (0.4-0.9 million years ago). AFLP analysis of 76 individuals of P. alpina, representing …

Long‐distance dispersal vs vicariance: the origin and genetic diversity of alpine plants in the Spanish Sierra Nevada

Here, we investigated the origin and genetic diversity of four alpine plant species co-occurring in the Spanish Sierra Nevada and other high mountains in south-western Europe by analysis of amplified fragment length polymorphisms (AFLPs). In Kernera saxatilis, Silene rupestris and Gentiana alpina we found intraspecific phylogroups corresponding to mountain regions as predicted by the vicariance hypothesis. Moreover, genetic distances between Sierra Nevada and Pyrenees populations were always higher than those between populations from the Pyrenees and the south-western Alps/Massif Central. This suggests successive disruption of gene exchange between mountain ranges as postglacial climatic wa…

Spatial and Temporal Patterns in the Evolution of the Flora of the European Alpine System

This paper presents a perspective of how phylogenetic and phylogeographic hypotheses, based on nuclear DNA sequence variation (ITS) or amplified fragment length polymorphisms (AFLPs), can provide insights into the origin and evolution of the European high mountain flora. We focus on a diversity of unrelated herbaceous plant taxa that are broadly co-distributed across the European Alpine System, representing different taxonomic levels, and having either Mediterranean or Asian affinities (i.e., Anthyllis montana, Pritzelago alpina, Globularia vs. Soldanella, and Primula sect. Auricula). Our observations highlight that all taxa investigated began to diversify at the beginning of the Pleistocen…