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The Afromontane and Afroalpine areas constitute some of the main biodiversity hotspots of Africa. They are particularly rich in plant endemics, but the biogeographic origins and evolutionary processes leading to this outstanding diversity are poorly understood. We performed phylogenomic and biogeographic analyses of one of the most species-rich plant genera in these mountains, Helichrysum (Compositae-Gnaphalieae). Most previous studies have focused on Afroalpine elements of Eurasian origin, and the southern African origin of Helichrysum provides an interesting counterexample. We obtained a comprehensive nuclear dataset from 304 species (≈50% of the genus) using target-enrichment with the Compositae1061 probe set. Summary-coalescent and concatenation approaches combined with paralog recovery yielded congruent, well-resolved phylogenies. Ancestral range estimations revealed that Helichrysum originated in arid southern Africa, whereas the southern African grasslands were the source of most lineages that dispersed within and outside Africa. Colonization of the tropical Afromontane and Afroalpine areas occurred repeatedly throughout the Miocene–Pliocene. This timing coincides with mountain uplift and the onset of glacial cycles, which together may have facilitated both speciation and intermountain gene flow, contributing to the evolution of the Afroalpine flora.

Alpine plant radiations are common across all major mountain systems of the world, and have been regarded as the main explanation for the species diversity found within these areas. To study the mechanisms behind the origin of this diversity, it is necessary to determine phylogenetic relationships and species boundaries in radiating alpine groups. The genus Dendrosenecio (Asteraceae) is an iconic example of a tropical-alpine plant radiation in the East African high mountains. To this date, limited sampling of molecular markers has resulted in insufficient phylogenetic resolution and infrageneric classification, hindering a comprehensive understanding of the drivers of diversification. Here, we used Hyb-Seq and the Compositae1061 probe set to generate targeted nuclear and off-target plastid DNA data for 42 samples representing all currently accepted 11 species. We combined coalescent methods and paralogy analysis to infer phylogenetic relationships, estimate divergence times and evaluate species boundaries. Lineage differentiation in Dendrosenecio seems to have occurred between the Late Miocene and the Pleistocene, starting when the first high elevation habitats became available in East Africa. We retrieved four major clades corresponding to four geographically distant mountain groups, testifying the importance of allopatric speciation in the early diversification of the group. Cytonuclear discordance suggested the occurrence of historical hybridization following occasional long-distance dispersal between mountain groups. The species delimitation analysis favored 10 species, but only five were fully supported, suggesting that population-level studies addressing processes such as ecological speciation and hybridization after secondary contact are needed to determine the current diversity found in the genus.

Summary ・Phytochromes play a central role in mediating adaptive responses to light and temperature throughout plant life cycles. Despite evidence for adaptive importance of natural variation in phytochromes, little information is known about molecular mechanisms that modulate physiological responses of phytochromes in nature. ・We show evolutionary divergence in physiological responses relevant to thermal stability of a physiologically active form of phytochrome (Pfr) between two sister species of Brassicaceae growing at different latitudes. The higher latitude species (Cardamine bellidifolia; Cb) responded more strongly to light‐limited conditions compared with its lower latitude sister (C. nipponica; Cn). Moreover, CbPHYB conferred stronger responses to both light‐limited and warm conditions in the phyB‐deficient mutant of Arabidopsis thaliana than CnPHYB: that is Pfr CbphyB was more stable in nuclei than CnphyB. ・Our findings suggest that fine tuning Pfr stability is a fundamental mechanism for plants to optimise phytochrome‐related traits in their evolution and adapt to spatially varying environments, and open a new avenue to understand molecular mechanisms that fine tune phytochrome responses in nature.

[Aim] The afroalpine sky-islands present one of the most interesting models to study discrete biogeographic patterns in a terrestrial island system. Here, we performed range-wide sampling of the afroalpine clade of fine-leaved Festuca grasses and address a set of hypotheses on its origin and dispersal. We focus on the widespread species F. abyssinica and explore the role of the eastern and western African sky-islands., [Location] Tropical Africa, [Taxon] Afroalpine Festuca, [Methods] We combine data from field surveys, phylogeography, coalescence-based dispersal modelling, and environmental niche and dispersal costs analyses to infer patterns of genetic diversity, genealogical relationships, colonization routes and range shifts under two Quaternary climates (current – to represent warm periods; and Last Glacial Maximum (LGM) – to represent cold periods)., [Results] The westernmost populations in East Africa show higher genetic diversity and higher similarities with the West African populations and the Ethiopian Simen Mountains than with the more closely situated East African populations. Dispersal models and ecological niche predictions of F. abyssinica supported multiple long-distance dispersals (LDD) among the eastern African sky-islands, and at least two dispersal events between the two sides of Africa (0.86 Ma and 0.52 Ma), probably facilitated by bridging suitable habitats during the coldest periods of the Pleistocene., [Main conclusions] We reconstruct an afroalpine mountain-hopping dispersal model, with migrations occurring between adjacent sky-islands in eastern Africa, and through a Central Africa–Sudan pathway connecting afroalpine patches on the two sides of the continent., The work was supported by the Spanish MINECO CGL2012-39953-C02-01 and the Norwegian NUFU 2007/1058 grant projects to PC and CB respectively. MN was funded by a Spanish Ministry of Education postdoctoral contract in the University of Zaragoza, and PC and MN by a Bioflora research team A01-17 and A01-20R grants co-funded by the Spanish Aragón Government and the European Social Fund (FEDER 2014-2020; "Construyendo Europa desde Aragón").

Tropical alpine floras are renowned for high endemism, spectacular giant rosette plants testifying to convergent adaptation to harsh climates with nightly frosts, and recruitment dominated by long-distance dispersal from remote areas. In contrast to the larger, more recent (late Miocene onward) and contiguous expanses of tropical alpine habitat in South America, the tropical alpine flora in Africa is extremely fragmented across small patches on distant mountains of variable age (Oligocene onward). How this has affected the colonization and diversification history of the highly endemic but species-poor afroalpine flora is not well known. Here we infer phylogenetic relationships of ∼20% of its species using novel genome skimming data and published matrices and infer a timeframe for species origins in the afroalpine region using fossil-calibrated molecular clocks. Although some of the mountains are old, and although stem node ages may substantially predate colonization, most lineages appear to have colonized the afroalpine during the last 5 or 10 My. The accumulation of species increased exponentially toward the present. Taken together with recent reports of extremely low intrapopulation genetic diversity and recent intermountain population divergence, this points to a young, unsaturated, and dynamic island scenario. Habitat disturbance caused by the Pleistocene climate oscillations likely induced cycles of colonization, speciation, extinction, and recolonization. This study contributes to our understanding of differences in the histories of recruitment on different tropical sky islands and on oceanic islands, providing insight into the general processes shaping their remarkable floras. acceptedVersion

The Arctic is one of the most extreme terrestrial environments on the planet. Here, we present the first chromosome‐scale genome assembly of a plant adapted to the high Arctic, Draba nivalis (Brassicaceae), an attractive model species for studying plant adaptation to the stresses imposed by this harsh environment. We used an iterative scaffolding strategy with data from short‐reads, single‐molecule long reads, proximity ligation data, and a genetic map to produce a 302 Mb assembly that is highly contiguous with 91.6% assembled into eight chromosomes (the base chromosome number). To identify candidate genes and gene families that may have facilitated adaptation to Arctic environmental stresses, we performed comparative genomic analyses with nine non‐Arctic Brassicaceae species. We show that the D. nivalis genome contains expanded suites of genes associated with drought and cold stress (e.g., related to the maintenance of oxidation‐reduction homeostasis, meiosis, and signaling pathways). The expansions of gene families associated with these functions appear to be driven in part by the activity of transposable elements. Tests of positive selection identify suites of candidate genes associated with meiosis and photoperiodism, as well as cold, drought, and oxidative stress responses. Our results reveal a multifaceted landscape of stress adaptation in the D. nivalis genome, offering avenues for the continued development of this species as an Arctic model plant., see also the Perspective by Tanja Pyhäjärvi and Tiina M. Mattila

Extreme environments offer powerful opportunities to study how different organisms have adapted to similar selection pressures at the molecular level. Arctic plants have adapted to some of the coldest and driest biomes on Earth and typically possess suites of similar morphological and physiological adaptations to extremes in light and temperature. Here, we compare patterns of molecular evolution in three Brassicaceae species that have independently colonized the Arctic and present some of the first genetic evidence for plant adaptations to the Arctic environment. By testing for positive selection and identifying convergent substitutions in orthologous gene alignments for a total of 15 Brassicaceae species, we find that positive selection has been acting on different genes, but similar functional pathways in the three Arctic lineages. The positively selected gene sets identified in the three Arctic species showed convergent functional profiles associated with extreme abiotic stress characteristic of the Arctic. However, there was little evidence for independently fixed mutations at the same sites and for positive selection acting on the same genes. The three species appear to have evolved similar suites of adaptations by modifying different components in similar stress response pathways, implying that there could be many genetic trajectories for adaptation to the Arctic environment. By identifying candidate genes and functional pathways potentially involved in Arctic adaptation, our results provide a framework for future studies aimed at testing for the existence of a functional syndrome of Arctic adaptation in the Brassicaceae and perhaps flowering plants in general.

Although logistically challenging to study, the Arctic is a bellwether for global change and is becoming a model for questions pertinent to the persistence of biodiversity. Disruption of Arctic ecosystems is accelerating, with impacts ranging from mixing of biotic communities to individual behavioral responses. Understanding these changes is crucial for conservation and sustainable economic development. Genomic approaches are providing transformative insights into biotic responses to environmental change, but have seen limited application in the Arctic due to a series of limitations. To meet the promise of genome analyses, we urge rigorous development of biorepositories from high latitudes to provide essential libraries to improve the conservation, monitoring, and management of Arctic ecosystems through genomic approaches.

Background and Aims The Arctic tundra, with its extreme temperatures and short growing season, is evolutionarily young and harbours one of the most species-poor floras on Earth. Arctic species often show little phenotypic and genetic divergence across circumpolar ranges. However, strong intraspecific post-zygotic reproductive isolation (RI) in terms of hybrid sterility has frequently evolved within selfing Arctic species of the genus Draba. Here we assess whether incipient biological species are common in the Arctic flora. Methods We conducted an extensive crossing experiment including six species representing four phylogenetically distant families collected across the circumpolar Arctic. We crossed conspecific parental populations representing different spatial scales, raised 740 F1 hybrids to maturity and measured fertility under laboratory conditions. We examined genetic divergence between populations for two of these species (Cardamine bellidifolia and Ranunculus pygmaeus). Key Results In five of the six species, we find extensive reduction in pollen fertility and seed set in F1 hybrids; 219 (46 %) of the 477 F1 hybrids generated between parents separated by ≥427 km had Conclusion We show that post-zygotic RI has developed multiple times within taxonomically recognized Arctic species belonging to several distantly related lineages, and that RI may have developed over just a few millennia. Rapid and widespread evolution of incipient biological species in the Arctic flora might be associated with frequent bottlenecks due to glacial cycles, and/or selfing mating systems, which are common in the harsh Arctic environment where pollinators are scarce.

The monumental work of Olov Hedberg provided deep insights into the spectacular and fragmented tropical alpine flora of the African sky islands. Here we review recent molecular and niche modelling studies and re-examine Hedberg’s hypotheses and conclusions. Colonisation started when mountain uplift established the harsh diurnal climate with nightly frosts, accelerated throughout the last 5 Myr (Plio-Pleistocene), and resulted in a flora rich in local endemics. Recruitment was dominated by long-distance dispersals (LDDs) from seasonally cold, remote areas, mainly in Eurasia. Colonisation was only rarely followed by substantial diversification. Instead, most of the larger genera and even species colonised the afroalpine habitat multiple times independently. Conspicuous parallel evolution occurred among mountains, e.g., of gigantism in Lobelia and Dendrosenecio and dwarf shrubs in Alchemilla. Although the alpine habitat was ~ 8 times larger and the treeline was ~ 1000 m lower than today during the Last Glacial Maximum, genetic data suggest that the flora was shaped by strong intermountain isolation interrupted by rare LDDs rather than ecological connectivity. The new evidence points to a much younger and more dynamic island scenario than envisioned by Hedberg: the afroalpine flora is unsaturated and fragile, it was repeatedly disrupted by the Pleistocene climate oscillations, and it harbours taxonomic and genetic diversity that is unique but severely depauperated by frequent bottlenecks and cycles of colonisation, extinction, and recolonisation. The level of intrapopulation genetic variation is alarmingly low, and many afroalpine species may be vulnerable to extinction because of climate warming and increasing human impact.

Distantly related lineages of the enigmatic giant rosette plants of tropical alpine environments provide classical examples of convergent adaptation. For the giant senecios (Dendrosenecio), the endemic landmarks of the East African sky islands, it has also been suggested that parallel adaptation has been important for within-lineage differentiation. To test this hypothesis and to address potential gene flow and hybridization among the isolated sky islands, we organized field expeditions to all major mountains. We sampled all currently accepted species and all but one subspecies and genotyped 460 plants representing 109 populations. We tested whether genetic structuring corresponds to geography, as predicted by a parallel adaptation hypothesis, or to altitudinal belt and habitat rather than mountains, as predicted by a hypothesis of a single origin of adaptations. Bayesian and Neighbor-Net analyses showed that the main genetic structure is shallow and largely corresponds to geography, supporting a hypothesis of recent, rapid radiation via parallel altitude/habitat adaptation on different mountains. We also found evidence for intermountain admixture, suggesting several long-distance dispersals by wind across vast areas of unsuitable habitat. The combination of parallel adaptation, secondary contact, and hybridization may explain the complex patterns of morphological variation and the contradicting taxonomic treatments of these rare enigmatic giants, supporting the use of wide taxonomic concepts. Notably, the within-population genetic diversity was very low and calls for increased conservation efforts.

DNA-based snapshots of ancient vegetation have shown that the composition of high-latitude plant communities changed considerably during the late Quaternary. However, parallel changes in biotic interactions remain largely uninvestigated. Here we show how mutualisms involving plants and heterotrophic organisms varied during the last 50,000 years. During 50–25 ka BP, a cool period featuring stadial-interstadial fluctuations, arbuscular mycorrhizal and non-N-fixing plants predominated. During 25-15 ka BP, a cold, dry interval, the representation of ectomycorrhizal, non-mycorrhizal and facultatively mycorrhizal plants increased, while that of N-fixing plants decreased further. From 15 ka BP, which marks the transition to and establishment of the Holocene interglaciation, representation of arbuscular mycorrhizal plants decreased further, while that of ectomycorrhizal, non-mycorrhizal, N-fixing and wind-pollinated plants increased. These changes in the mutualist trait structure of vegetation may reflect responses to historical environmental conditions that are without current analogue, or biogeographic processes, such as spatial decoupling of mutualist partners., Recently, an eDNA metabarcoding data set was used to describe northern high-latitude vegetation during the past 50,000 years. Here, Zobel et al. use the data set to examine how the abundance of key plant mutualistic traits changed during this period and discuss possible environmental drivers.

Compilation of the species richness data was made possible through the TFI Networks grant to CD, “Effect Studies and Adaptation to Climate Change,” under the Norforsk initiative (2011 – 2014) which supported two CBIONET-AVA workshops held in Denmark during 2013. The genetic studies were funded by the Research Council of Norway (grant nos. 150322/720 and 170952/V40 to CB).

Published version available at https://doi.org/10.1007/s00035-015-0162-2. The naturally fragmented habitat on the tallest African mountains provides a good model system to study vicariance, dispersal, and hybridization. Many mountains are separated by lowland that likely was unsuitable for high-alpine plants even during cold climatic periods. We explore the relative importance of these processes using two endemic sister species: the widespread Ethiopian/eastern East African Carex monostachya and the mainly western East African C. runssoroensis. These bog-forming sedges co-occur in some mountains and are hypothesized to hybridize. The two species were distinctly differentiated for genome-wide Amplified Fragment Length Polymorphisms (AFLPs), also in one mountain where they co-occur. However, the plants from another mountain showed strong signals of admixture. The results suggest initial divergence into one western and one northern/eastern lineage, followed by long-distance dispersal resulting in secondary contact zones. Also within species genetic diversity was clearly structured with distinct genetic groups on some, but not all mountains. Differentiation levels varied considerably and did not always correspond to the extent of lowland habitat between mountains. The narrow Rift Valley in the otherwise nearly contiguous highlands in Ethiopia appears to present a much stronger barrier to dispersal than the extensive lowlands separating Ethiopia from East Africa. This may be a general pattern since it has been documented also for other afro-alpine species

Premise of the study Many arctic-alpine species have vast geographic ranges, but these may encompass substantial gaps whose origins are poorly understood. Here we address the phylogeographic history of Silene acaulis, a perennial cushion plant with a circumpolar distribution except for a large gap in Siberia. Methods We assessed genetic variation in a range-wide sample of 103 populations using plastid DNA (pDNA) sequences and AFLPs (amplified fragment length polymorphisms). We constructed a haplotype network and performed Bayesian phylogenetic analyses based on plastid sequences. We visualized AFLP patterns using principal coordinate analysis, identified genetic groups using the program structure, and estimated genetic diversity and rarity indices by geographic region. Key results The history of the main pDNA lineages was estimated to span several glaciations. AFLP data revealed a distinct division between Beringia/North America and Europe/East Greenland. These two regions shared only one of 17 pDNA haplotypes. Populations on opposite sides of the Siberian range gap (Ural Mountains and Chukotka) were genetically distinct and appear to have resulted from postglacial leading-edge colonizations. We inferred two refugia in North America (Beringia and the southern Rocky Mountains) and two in Europe (central-southern Europe and northern Europe/East Greenland). Patterns in the East Atlantic region suggested transoceanic long-distance dispersal events. Conclusions Silene acaulis has a highly dynamic history characterized by vicariance, regional extinction, and recolonization, with persistence in at least four refugia. Long-distance dispersal explains patterns across the Atlantic Ocean, but we found no evidence of dispersal across the Siberian range gap.

Birches (Betula spp.) hybridize readily, confounding genetic signatures of refugial isolation and postglacial migration. We aimed to distinguish hybridization from range-shift processes in the two widespread and cold-adapted species B. nana and B. pubescens, previously shown to share a similarly east-west-structured variation in plastid DNA (pDNA). We sampled the two species throughout their ranges and included reference samples of five other Betula species and putative hybrids. We analysed 901 individual plants using mainly nuclear high-resolution markers (amplified fragment length polymorphisms; AFLPs); a subset of 64 plants was also sequenced for two pDNA regions. Whereas the pDNA variation as expected was largely shared between B. nana and B. pubescens, the two species were distinctly differentiated at AFLP loci. In B. nana, both the AFLP and pDNA results corroborated the former pDNA-based hypothesis that it expanded from at least two major refugia in Eurasia, one south of and one east of the North European ice sheets. In contrast, B. pubescens showed a striking lack of geographic structuring of its AFLP variation. We identified a weak but significant increase in nuclear (AFLP) gene flow from B. nana into B. pubescens with increasing latitude, suggesting hybridization has been most frequent at the postglacial expansion front of B. pubescens and that hybrids mainly backcrossed to B. pubescens. Incongruence between pDNA and AFLP variation in B. pubescens can be explained by efficient expansion from a single large refugium combined with leading-edge hybridization and plastid capture from B. nana during colonization of new territory already occupied by this more cold-tolerant species.

High Arctic environments are particularly sensitive to climate changes, but retrieval of paleoecological data is challenging due to low productivity and biomass. At the same time, Arctic soils and sediments have proven exceptional for long-term DNA presentation due to their constantly low temperatures. Lake sediments contain DNA paleorecords of the surrounding ecosystems and can be used to retrieve a variety of organismal groups from a single sample. In this study, we analyzed vascular plant, bryophyte, algal (in particular diatom) and copepod DNA retrieved from a sediment core spanning the Holocene, taken from Bliss Lake on the northernmost coast of Greenland. A previous multi-proxy study including microscopic diatom analyses showed that this lake experienced changes between marine and lacustrine conditions. We inferred the same environmental changes from algal DNA preserved in the sediment core. Our DNA record was stratigraphically coherent, with no indication of leaching between layers, and our cross-taxon comparisons were in accordance with previously inferred local ecosystem changes. Authentic ancient plant DNA was retrieved from nearly all layers, both from the marine and the limnic phases, and distinct temporal changes in plant presence were recovered. The plant DNA was mostly in agreement with expected vegetation history, but very early occurrences of vascular plants, including the woody Empetrum nigrum, document terrestrial vegetation very shortly after glacial retreat. Our study shows that multi-taxon metabarcoding of sedimentary ancient DNA from lake cores is a valuable tool both for terrestrial and aquatic paleoecology, even in low-productivity ecosystems such as the High Arctic. (C) 2015 Elsevier Ltd. All rights reserved.

Arctic-alpine plants have expanded and contracted their ranges in response to the Pleistocene climate oscillations. Today, many arctic-alpine plants have vast distributions in the circumarctic region as well as marginal, isolated occurrences in high mountains at lower latitudes. These marginal populations may represent relict, long-standing populations that have persisted for several cycles of cold and warm climate during the Pleistocene, or recent occurrences that either result from southward step-wise migration during the last glacial period or from recent long-distance dispersal. In light of these hypotheses, we investigated the biogeographic history of the marginal Japanese populations of the widespread arctic-alpine plant Vaccinium vitis-idaea (Ericaceae), which is bird-dispersed, potentially over long distances. We sequenced three nuclear loci and one plastid DNA region in 130 individuals from 65 localities covering its entire geographic range, with a focus on its marginal populations in Japan. We found a homogenous genetic pattern across its enormous range based on the loci analysed, in contrast to the geographically structured variation found in a previous study of amplified fragment length polymorphisms in this species. However, we found several unique haplotypes in the Japanese populations, excluding the possibility that these marginal populations result from recent southward migration. Thus, even though V. vitis-idaea is efficiently dispersed via berries, our study suggests that its isolated populations in Japan have persisted during several cycles of cold and warm climate during the Pleistocene.

Aim We used combined palaeobotanical and genetic data to assess whether Norway spruce (Picea abies) and Siberian spruce (Picea obovata), two major components of the Eurasian boreal forests, occupied separate glacial refugia, and to test previous hypotheses on their distinction, geographical delimitation and introgression. Location The range of Norway spruce in northern Europe and Siberian spruce in northern Asia. Methods Pollen data and recently compiled macrofossil records were summarized for the Last Glacial Maximum (LGM), late glacial and Holocene. Genetic variation was assessed in 50 populations using one maternally (mitochondrial nad1) and one paternally (chloroplast trnT–trnL) inherited marker and analysed using spatial analyses of molecular variance (SAMOVA). Results Macrofossils showed that spruce was present in both northern Europe and Siberia at the LGM. Congruent macrofossil and pollen data from the late glacial suggested widespread expansions of spruce in the East European Plain, West Siberian Plain, southern Siberian mountains and the Baikal region. Colonization was largely completed during the early Holocene, except in the formerly glaciated area of northern Europe. Both DNA markers distinguished two highly differentiated groups that correspond to Norway spruce and Siberian spruce and coincide spatially with separate LGM spruce occurrences. The division of the mtDNA variation was geographically well defined and occurred to the east of the Ural Mountains along the Ob River, whereas the cpDNA variation showed widespread admixture. Genetic diversity of both DNA markers was higher in western than in eastern populations. Main conclusions North Eurasian Norway spruce and Siberian spruce are genetically distinct and occupied separate LGM refugia, Norway spruce on the East European Plain and Siberian spruce in southern Siberia, where they were already widespread during the late glacial. They came into contact in the basin of the Ob River and probably hybridized. The lower genetic diversity in the eastern populations may indicate that Siberian spruce suffered more from past climatic fluctuations than Norway spruce.

Aim Successful colonization after long-distance dispersal (LDD) depends on the availability of suitable habitats and competitive ability. In this study, we address the hypothesis that two widely distributed sister grass lineages (broad- and fine-leaved Loliinae; hereafter BL and FL) that differ in their habitat requirements and competitive ability also differ in their biogeographic history, with particular reference to LDD. Location Global. Methods We reconstructed a comprehensive phylogeny of Loliinae, based on nuclear and plastid markers and estimated divergence times using fossil calibrations. Biogeographical events were estimated using analysis of range evolution, comparing different models. Numbers and rates of dispersals were estimated for BL and FL using stochastic mapping with best-performing and baseline biogeographical models, and examined for correlation with distance, disjunction type, and phenotypic syndrome. Results The most recent common ancestor of Loliinae likely split at the Oligocene-Miocene transition (22.50 ± 3.95 Ma), pre-dating previous estimates, whereas the ancestors of the BL and FL Loliinae likely began to diversify in the Early Miocene (18.91 ± 4.15 and 17.50 ± 3.50 Ma, respectively). A model of range evolution integrating founder events and scaling of dispersal by shortest distance between areas performed best amongst a set of alternative models and recovered a mean of 83 dispersal events in Loliinae. Overall dispersal rates were significantly higher in BL than in FL. Per-route dispersal rates showed a significant negative exponential relationship to shortest distance but were not affected by phenotypic syndrome or disjunction type. Main conclusions Loliinae originated in the Northern Hemisphere and evolved through recurrent LDDs. Higher competitive ability, potentially related to the broad-leaved syndrome (i.e. tall strong-rhizomatous plants, long-living individuals, occupancy of more stable habitats), may explain higher observed dispersal rates in BL compared with FL Loliinae. However, the dominant factor impacting dispersal in both BL and FL Loliinae is the distance between suitable areas.

The tropical alpine ecosystem in eastern Africa is highly fragmented among biological ‘sky islands’, where populations of frost-tolerant organisms are isolated from each other by a ‘sea’ of tropical lowlands. One-third of the species in the afroalpine flora are exclusively alpine, but the other species can to varying degrees extend into grasslands and open forests of lower vegetation belts. A long-debated question is whether colonization of the alpine zone of these mountains and subsequent intermountain gene flow entirely depend on long-distance dispersal across unsuitable habitats, or whether suitable habitats shifted far enough downslope under past colder climates to form bridges enabling gradual migration. Here we address this question using a classification tree model. We mapped the extent of the current alpine habitat and projected it to the last glacial maximum (LGM) climate to assess whether gradual migration was possible for exclusively alpine taxa during this glacial period, and thus potentially also during earlier Pleistocene glaciations. Next, we modelled landcover under current and LGM climates to assess whether grassland and open forests could have served as migration corridors for alpine taxa that today extend into lower vegetation belts. We estimated that the LGM treeline was about 1000 m lower and the alpine habitat was about eight times larger than that today. At the LGM, we found that most of the currently fragmented alpine habitat of the Ethiopian highlands was interconnected except across the Great Rift Valley, whereas the solitary mountains of East/Central Africa remained isolated for exclusively alpine species. However, for drought-tolerant alpine species that today extend below the treeline, gradual migration through habitat corridors may have been possible among mountains during the dry glacial periods, and possibly also under the current climate before agriculture transformed the low-lying landscapes. Afroalpine · CART · Gene flow · Habitat connectivity · Last glacial maximum · Treeline This is a post-peer-review, pre-copyedit version of an article published in Alpine Botany. The final authenticated version is available online at: http://dx.doi.org/10.1007/s00035-017-0184-z

The scattered eastern African high mountains harbor a renowned and highly endemic flora, but the taxonomy and phylogeographic history of many plant groups are still insufficiently known. The high-alpine populations of the Geranium arabicum/kilimandscharicum complex present intricate morphological variation and have recently been suggested to comprise two new endemic taxa. Here we aim to contribute to a clarification of the taxonomy of these populations by analyzing genetic (AFLP) variation in range-wide high-alpine samples, and we address whether hybridization has contributed to taxonomic problems. We identified only two genetic groups. One corresponded to G. kilimandscharicum, which has been reported as exclusively high-alpine and confined to the eastern Rift mountains in East Africa. The other corresponded to G. arabicum, reported from lower altitudes on the same mountains as well as from a wide altitudinal span in Ethiopia and on the western Rift mountains in East Africa. The four populations analyzed of a recently described species from the Bale Mts in Ethiopia were admixed, indicating that they result from recent long-distance dispersal of G. kilimandscharicum from East Africa followed by hybridization with local G. arabicum in naturally disturbed habitats. Some admixture between the two genetic groups was also inferred on other mountains, supporting earlier suggestions of introgression based on morphology. We did not find support for recognition of the recently suggested new subspecies of G. arabicum in Ethiopia. Interestingly, the high-alpine G. kilimandscharicum lacked clear geographic structuring, suggesting a recent history of colonization of the different mountains or extensive intermountain gene flow.

International audience; Although it is generally agreed that the Arctic flora is among the youngest and least diverse on Earth, the processes that shaped it are poorly understood. Here we present 50 thousand years (kyr) of Arctic vegetation history, derived from the first large-scale ancient DNA metabarcoding study of circumpolar plant diversity. For this interval we also explore nematode diversity as a proxy for modelling vegetation cover and soil quality, and diets of herbivorous megafaunal mammals, many of which became extinct around 10 kyr bp (before present). For much of the period investigated, Arctic vegetation consisted of dry steppe-tundra dominated by forbs (non-graminoid herbaceous vascular plants). During the Last Glacial Maximum (25-15 kyr bp), diversity declined markedly, although forbs remained dominant. Much changed after 10 kyr bp, with the appearance of moist tundra dominated by woody plants and graminoids. Our analyses indicate that both graminoids and forbs would have featured in megafaunal diets. As such, our findings question the predominance of a Late Quaternary graminoid-dominated Arctic mammoth steppe.

Conservation of biodiversity may in the future increasingly depend upon the availability of scientific information to set suitable restoration targets. In traditional paleoecology, sediment-based pollen provides a means to define preanthropogenic impact conditions, but problems in establishing the exact provenance and ecologically meaningful levels of taxonomic resolution of the evidence are limiting. We explored the extent to which the use of sedimentary ancient DNA (sedaDNA) may complement pollen data in reconstructing past alpine environments in the tropics. We constructed a record of afro-alpine plants retrieved from DNA preserved in sediment cores from 2 volcanic crater sites in the Albertine Rift, eastern Africa. The record extended well beyond the onset of substantial anthropogenic effects on tropical mountains. To ensure high-quality taxonomic inference from the sedaDNA sequences, we built an extensive DNA reference library covering the majority of the afro-alpine flora, by sequencing DNA from taxonomically verified specimens. Comparisons with pollen records from the same sediment cores showed that plant diversity recovered with sedaDNA improved vegetation reconstructions based on pollen records by revealing both additional taxa and providing increased taxonomic resolution. Furthermore, combining the 2 measures assisted in distinguishing vegetation change at different geographic scales; sedaDNA almost exclusively reflects local vegetation, whereas pollen can potentially originate from a wide area that in highlands in particular can span several ecozones. Our results suggest that sedaDNA may provide information on restoration targets and the nature and magnitude of human-induced environmental changes, including in high conservation priority, biodiversity hotspots, where understanding of preanthropogenic impact (or reference) conditions is highly limited.

The flora on the afro-alpine sky islands is renowned for extreme fragmentation, representing a unique natural experiment in biogeography. Here we address the roles of isolation and gene flow, in particular across the narrow Rift Valley (the RV barrier) that cuts through the Ethiopian Highlands (EH), and across the vast low-lying landscape that separates EH from the East African mountains (the EH–EA barrier). We inferred the history of two species with different dispersal mechanisms, but with similar geographic ranges and habitats based on Amplified fragment length polymorphisms (AFLPs). Contrary to our predictions, we found that the populations from opposite sides of the RV barrier were less similar than those from opposite sides of the EH–EA barrier, and that only the supposedly short distance-dispersed species (Trifolium cryptopodium) showed a strong signal of secondary gene flow across the RV barrier. In the wind-dispersed Carduus schimperi, we rather found an evidence for the gene flow between differentiated populations inhabiting different EA mountains. Both species harbored little genetic diversity but considerable genetic rarity in several individual mountains, suggesting long-term isolation and bottlenecks during climatically unfavorable periods. Our genetic data corroborate a division of C. schimperi into three subspecies, but with new delimitation of their ranges, and of T. cryptopodium into two intraspecific taxa. Our findings support the idea that stochasticity may play a major role in shaping extremely fragmented ecosystems such as the afro-alpine. After initial colonization of different mountains, periods of isolation may alternate with unpredictable episodes of intermountain gene flow.

Background and aims Repeated hybridization and/or polyploidization confound classification and phylogenetic inference, and multiple colonizations at different time scales complicate biogeographical reconstructions. This study investigates whether such processes can explain long-term controversies in Anthoxanthum, and in particular its debated relationship to the genus Hierochloe, the evolution of its conspicuously diverse floral morphology, and the origins of its strikingly disjunct occurrences. A hypothesis for recurrent polyploid formation is proposed. Methods Three plastid (trnH-psbA, trnT-L and trnL-F) and two nuclear (ITS, ETS) DNA regions were sequenced in 57 accessions of 17 taxa (including 161 ETS clones) and Bayesian phylogenetic analyses were conducted. Divergence times were inferred in *BEAST using a strict molecular clock. Key results Anthoxanthum was inferred as monophyletic and sister to one species of Hierochloe based on the plastid data, whereas the nuclear data suggested that one section (Anthoxanthum section Anthoxanthum) is sister to a clade including the other section (Anthoxanthum section Ataxia) as sister to the genus Hierochloe. This could explain the variation in floral morphology; the aberrant characters in Ataxia seem to result from a Miocene hybridization event between one lineage with one fertile and two sterile florets (the Anthoxanthum lineage) and one which probably had three fertile florets as in extant Hierochloe. The distinct diploid A. gracile lineage originated in the Miocene; all other speciation events, many of them involving polyploidy, were dated to the Late Pliocene to Late Pleistocene. Africa was apparently colonized twice in the Late Pliocene (from the north to afro-alpine eastern Africa, and from south-east Asia to southern Africa), whereas Macaronesia was colonized much later (Late Pleistocene) by a diploid Mediterranean lineage. The widespread European tetraploid A. odoratum originated at least twice. Conclusions Many of the controversies in Anthoxanthum can be explained by recurring hybridization and/or polyploidization on time scales ranging from the Miocene to the Late Pleistocene. All but one of the extant species shared most recent common ancestors in the Late Pliocene to the Late Pleistocene. The disjunct occurrences in Africa originated in the Late Pliocene via independent immigrations, whereas Macaronesia was colonized in the Late Pleistocene.

The ericaceous vegetation zone of the unique and highly fragmented afro-alpine environment in the eastern African high mountains is typically dominated by the heather Erica arborea , often in combination with its close relative E. trimera. Both species are shrubs or small trees with tiny seeds, potentially capable of dispersal by wind over long distances. While E. arborea is widely distributed in Africa, the Middle East and Europe, E. trimera is endemic to the afro-alpine region where it is restricted to higher altitudes than E. arborea . We used Amplified Fragment Length Polymorphisms (AFLPs) and variation in non-coding plastid DNA sequences to test whether these two morphologically and ecologically very similar species display similar phylogeographic patterns in the afro-alpine region. We predict that the more high-altitudinal E. trimera shows more distinct genetic structuring than E. arborea , because dispersal of the latter may have been facilitated by formation of interglacial forest bridges between mountains. Based on extensive field sampling in most of the high mountains of Ethiopia and East Africa, we show that the two species are clearly distinct at AFLP and plastid DNA loci. Both showed low levels of overall AFLP diversity, suggesting bottlenecking in small refugial populations during unfavourable climatic periods. However, their genetic structuring and inferred phylogeographic histories were conspicuously different. The more high-altitudinal E. trimera consisted of three to four distinct AFLP groups, which also had different plastid DNA haplotypes and different geographic distributions, suggesting long-term restriction to several refugia (at least one in Ethiopia and two in East Africa). In contrast, E. arborea showed little geographic structuring at AFLP loci and only a single, widespread plastid DNA haplotype, which may suggest recent colonization of the entire study area from a single source population, likely via a combination of gradual expansion via forest bridges and long-distance dispersals. The source population of E. arborea may be situated in (or north of) Ethiopia, which harbours most genetic diversity.

High tropical mountains harbour remarkable and fragmented biodiversity thought to a large degree to have been shaped by multiple dispersals of cold-adapted lineages from remote areas. Few dated phylogenetic/phylogeographic analyses are however available. Here, we address the hypotheses that the sub-Saharan African sweet vernal grasses have a dual colonization history and that lineages of independent origins have established secondary contact. We carried out rangewide sampling across the eastern African high mountains, inferred dated phylogenies from nuclear ribosomal and plastid DNA using Bayesian methods, and performed flow cytometry and AFLP (amplified fragment length polymorphism) analyses. We inferred a single Late Pliocene western Eurasian origin of the eastern African taxa, whose high-ploid populations in one mountain group formed a distinct phylogeographic group and carried plastids that diverged from those of the currently allopatric southern African lineage in the Mid- to Late Pleistocene. We show that Anthoxanthum has an intriguing history in sub-Saharan Africa, including Late Pliocene colonization from southeast and north, followed by secondary contact, hybridization, allopolyploidization and local extinction during one of the last glacial cycles. Our results add to a growing body of evidence showing that isolated tropical high mountain habitats have a dynamic recent history involving niche conservatism and recruitment from remote sources, repeated dispersals, diversification, hybridization and local extinction. The final version of this research has been published in Molecular Ecology. © 2017 Wiley

The circumarctic ranges of arctic-alpine plants are thought to have been established in the late Pliocene/early Pleistocene, when the modern arctic tundra was formed in response to climate cooling. Previous findings of range-wide genetic structure in arctic-alpine plants have been thought to support this hypothesis, but few studies have explicitly addressed the temporal framework of the genetic structure. Here, we estimated the demographic history of the genetic structure in the circumarctic Kalmia procumbens using sequences of multiple nuclear loci and examined whether its genetic structure reflects prolonged isolation throughout the Pleistocene. Both Bayesian clustering and phylogenetic analyses revealed genetic distinction between alpine and arctic regions, whereas detailed groupings were somewhat discordant between the analyses. By assuming a population grouping based on the phylogenetic analyses, which likely reflects a deeper intraspecific divergence, we conducted model-based analyses and demonstrated that the intraspecific genetic divergence in K. procumbens likely originated during the last glacial period. Thus, there is no need to postulate range separation throughout the Pleistocene to explain the current genetic structure in this species. This study demonstrates that range-wide genetic structure in arctic-alpine plants does not necessarily result from the late Pliocene/early Pleistocene origin of their circumarctic ranges, and emphasizes the importance of a temporal framework of the current genetic structure for understanding the biogeographic history of the arctic flora. This article is protected by copyright. All rights reserved.

Aim High intra specific genetic diversity is necessary for species adaptation to novel environments under climate change but species tracking suitable conditions are losing alleles through successive founder events during range shift. Here we investigated the relationship between range shift since the Last Glacial Maximum (LGM) and extant population genetic diversity across multiple plant species to understand variability in species responses. Location The circumpolar Arctic and northern temperate alpine ranges. Methods We estimated the climatic niches of 30 cold adapted plant species using range maps coupled with species distribution models and hindcasted species suitable areas to reconstructions of the mid Holocene and LGM climates. We computed the species specific migration distances from the species glacial refugia to their current distribution and correlated distances to extant genetic diversity in 1295 populations. Differential responses among species were related to life history traits. Results We found a negative association between inferred migration distances from refugia and genetic diversities in 25 species but only 11 had statistically significant negative slopes. The relationships between inferred distance and population genetic diversity were steeper for insect pollinated species than wind pollinated species but the difference among pollination system was marginally independent from phylogenetic autocorrelation. Main conclusion The relationships between inferred migration distances and genetic diversities in 11 species independent from current isolation indicate that past range shifts were associated with a genetic bottleneck effect with an average of 21 loss of genetic diversity per 1000 km-1. In contrast the absence of relationship in many species also indicates that the response is species specific and may be modulated by plant pollination strategies or result from more complex historical contingencies than those modelled here.

This article is also published at http://doi.wiley.com/10.1002/ece3.2603 This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. The main aim of this paper is to address consequences of climate warming on loss of habitat and genetic diversity in the enigmatic tropical alpine giant rosette plants using the Ethiopian endemic Lobelia rhynchopetalum as a model. We modeled the habitat suitability of L. rhynchopetalum and assessed how its range is affected under two climate models and four emission scenarios. We used three statistical algorithms calibrated to represent two different complexity levels of the response. We analyzed genetic diversity using amplified fragment length polymorphisms and assessed the impact of the projected range loss. Under all model and scenario combinations and consistent across algorithms and complexity levels, this afro-alpine flagship species faces massive range reduction. Only 3.4% of its habitat seems to remain suitable on average by 2,080, resulting in loss of 82% (CI 75%–87%) of its genetic diversity. The remaining suitable habitat is projected to be fragmented among and reduced to four mountain peaks, further deteriorating the probability of long-term sustainability of viable populations. Because of the similar morphological and physiological traits developed through convergent evolution by tropical alpine giant rosette plants in response to diurnal freeze-thaw cycles, they most likely respond to climate change in a similar way as our study species. We conclude that specialized high-alpine giant rosette plants, such as L. rhynchopetalum, are likely to face very high risk of extinction following climate warming

Human population expansion and associated degradation of the habitat of many wildlife species cause loss of biodiversity and species extinctions. The small Simen Mountains National Park in Ethiopia is one of the last strongholds for the preservation of a number of afro-alpine mammals, plants and birds, and it is home to the rare endemic Walia ibex, Capra walie. The narrow distribution range of this species as well as potential competition for resources with livestock, especially with domestic goat, Capra hircus, may compromise its future survival. Based on a curated afro-alpine taxonomic reference library constructed for plant taxon identification, we investigated the diet of the Walia ibex and addressed the dietary overlap with domestic goat using DNA metabarcoding of faecal samples. Faeces of both species were collected from different localities in the National Park. We show that both species are browsers, with forbs, shrubs and trees comprising the largest proportion of their diet, supplemented by grasses. There was a considerable overlap in dietary preferences. Several of the preferred diet items of the Walia ibex (Alchemilla sp., Hypericum revolutum, Erica arborea and Rumex sp.) were also among the most preferred diet items of the domestic goat. These results indicate that there is potential for competition between the two species, especially during the dry season, when resources are limited. Our findings, in combination with the expected increase in domestic herbivores, suggest that management plans should consider the potential threat posed by domestic goats to ensure future survival of the endangered Walia ibex. published

Aim Disjunctly distributed peatmosses (Sphagnum) have been found to exhibit little genetic structure over regional and intercontinental scales, mainly caused by high ability for transoceanic long-distance dispersal. Although, most Northern Hemisphere peatmoss species have wide circumboreal/nemoral ranges, little is known about the magnitude and effects of long-distance dispersal and barriers in shaping the genetic structure of such species. We investigate whether high dispersal capacity has caused genetic homogeneity across broad areas of the Northern Hemisphere, or whether barriers act to shape genetic structure across different species with similar distributional ranges. Location Northern Hemisphere. Methods We studied genetic variation and structure in six Sphagnum species using 19 microsatellite loci. Results Four out of six species were genetically structured in similar ways; with mainly one Beringian and one Atlantic group. Overall, both the North American and Eurasian continents seemed to act as a barrier to gene flow in several species. However, the most abrupt breakpoint between genetic groups was found in south-east Alaska. Main conclusions We found evidence for extensive gene flow between regions across the Northern Hemisphere among peatmosses, with oceans seemingly acting as weaker barriers to gene flow than landmasses. Plants from the amphi-Atlantic and amphi-Beringian regions of several species were genetically differentiated. Similar genetic structuring across several species, indicate that spore-producing species do not disperse freely across their entire distributional range, but are likely limited by wind directions, landmass barriers and/or habitat availability. This is the peer reviewed version of the following article: [Long-distance dispersal and barriers shape genetic structure of peatmosses (Sphagnum) across the Northern Hemisphere], which has been published in final form at [http://onlinelibrary.wiley.com/doi/10.1111/jbi.12716/abstract]. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving.

The flora on the isolated high African mountains or ‘sky islands’ is remarkable for its peculiar adaptations, local endemism and striking biogeographical connections to remote parts of the world. Ages of the plant lineages and the timing of their radiations have frequently been debated but remain contentious as there are few estimates based on explicit models and fossil-calibrated molecular clocks. We used the plastid region maturaseK (matK) and a Caryophylloflora paleogenica fossil to infer the age of the genus Lychnis, and constructed a data set of three plastid (matK; a ribosomal protein S16 (rps16); and an intergenic spacer (psbE-petL)) and two nuclear (internal transcribed spacer (ITS) and a region spanning exon 18-24 in the second largest subunit of RNA polymerase II (RPB2)) loci for joint estimation of the species tree and divergence time of the African representatives. The time of divergence of the African high-altitude Lychnis was placed in the late Miocene to early Pliocene. A single speciation event was inferred in the early Pliocene; subsequent speciation took place sporadically from the late Pliocene to the middle Pleistocene. We provide further support for a Eurasian origin of the African ‘sky islands’ floral elements, which seem to have been recruited via dispersals at different times: some old, as in Lychnis, and others very recent. We show that dispersal and diversification within Africa play an important role in shaping these isolated plant communities. This is the pre-peer reviewed version of the article, which has been published in final form at https://doi.org/10.1111/nph.13937. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving."

The taxonomic and ecological diversity of ancient fungal communities was assessed by combining next generation sequencing and metabarcoding of DNA preserved in permafrost. Twenty-six sediment samples dated 16 000-32 000 radiocarbon years old from two localities in Siberia were analysed for fungal ITS. We detected 75 fungal OTUs from 21 orders representing three phyla, although rarefaction analyses suggested that the full diversity was not recovered despite generating an average of 6677 ± 3811 (mean ± SD) sequences per sample and that preservation bias likely has considerable effect on the recovered DNA. Most OTUs (75.4%) represented ascomycetes. Due to insufficient sequencing depth, DNA degradation and putative preservation biases in our samples, the recovered taxa probably do not represent the complete historic fungal community, and it is difficult to determine whether the fungal communities varied geographically or experienced a composition shift within the period of 16 000-32 000 bp. However, annotation of OTUs to functional ecological groups provided a wealth of information on the historic communities. About one-third of the OTUs are presumed plant-associates (pathogens, saprotrophs and endophytes) typical of graminoid- and forb-rich habitats. We also detected putative insect pathogens, coprophiles and keratinophiles likely associated with ancient insect and herbivore faunas. The detection of putative insect pathogens, mycoparasites, aquatic fungi and endophytes broadens our previous knowledge of the diversity of fungi present in Beringian palaeoecosystems. A large group of putatively psychrophilic/psychrotolerant fungi was also detected, most likely representing a modern, metabolically active fungal community.

Aim Arctic plant phylogeography has largely focused on seed plants, and studies on other plant groups are necessary for comparison. Bryophytes have a unique life cycle and can be resistant to extreme conditions, suggesting that their phylogeographic patterns may differ from those of vascular plants. We address the history of the bryophyte genus Cinclidium in order to assess: (1) interspecific relationships, (2) whether its current broad circumarctic distribution results from recent dispersal or has been maintained by long-term local survival under severe glacial conditions, and (3) the origin of its bipolar disjunction. Location Arctic/boreal and bipolar. Methods We sequenced three plastid regions (atpH–atpI, rpl32–trnL and clpP1.1–clpP1.2) in 129 accessions covering the entire geographical range of all four described species, and inferred phylogenetic relationships and phylogeographical patterns using maximum parsimony, statistical parsimony and Bayesian inference. Results Cinclidium subrotundum was inferred to be monophyletic, in agreement with its distinct morphology and ecology. The three remaining known species (the haploids C. latifolium and C. arcticum, and the diploid C. stygium) shared a number of closely related or identical haplotypes despite their clear morphological differentiation. In all species, identical haplotypes occurred across the entire circumpolar region, including North Atlantic islands. In the bipolar species C. stygium, the haplotype observed in South America (Tierra del Fuego) was identical to one found in Iceland. Three populations originally referred to C. latifolium harboured highly divergent haplotypes and may represent a new species. Main conclusions The extensive haplotype sharing suggests a polyploid origin of C. stygium from C. arcticum, as well as incomplete lineage sorting and/or hybridization between the two haploids C. arcticum and C. latifolium. We interpret the finding of identical haplotypes over vast areas, including isolated islands, as a result of recent dispersal causing the circumpolar distribution of all species in the Northern Hemisphere and the extreme bipolar disjunction in C. stygium. The patterns in the bryophyte genus Cinclidium resemble those previously documented in arctic–alpine and bipolar vascular plants, suggesting that similar mechanisms have shaped species distributions in bryophytes and higher plants.

Aim Based on extensive range-wide sampling, we address the phylogeographicalhistory of one of the most widespread and taxonomically complex sedges inEurope, Carex nigra s. lat. We compare the genetic structure of the recentlycolonized northern areas (front edge) and the long-standing southern areas (rearedge), and assess the potential genetic basis of suggested taxonomic divisions atthe rank of species and below.Location Amphi-Atlantic, central and northern Europe, circum-Mediterraneanmountain ranges, central Siberia, Himalayas.Methods A total of 469 individuals sampled from 83 populations, covering mostof the species’ range, were analysed with amplified fragment lengthpolymorphism (AFLP) and chloroplast DNA (cpDNA) markers. Bayesianclustering, principal coordinates analysis, and estimates of diversity anddifferentiation were used for the analysis of AFLP data. CpDNA data wereanalysed with statistical parsimony networks and maximum parsimony andBayesian inference of phylogenetic trees.Results Overall genetic diversity was high, but differentiation among populationswas limited. Major glacial refugia were inferred in the Mediterranean Basin and inwestern Russia; in addition, there may have been minor refugia in the NorthAtlantic region. In the southern part of the range, we found high levels, butgeographically quite poorly structured genetic diversity, whereas the levels ofgenetic diversity varied among different areas in the north. North Americanpopulations were genetically very similar to the European populations.Main conclusions The data are consistent with extensive gene flow, which hasobscured the recent history of the taxon. The limited differentiation in the southprobably results from the mixing of lineages expanding from several local refugia.Northward post-glacial colonization resulted in a leading-edge pattern of lowdiversity in the Netherlands, Belgium, Scotland and Iceland, whereas the observedhigh diversity levels in Fennoscandia suggest broad-fronted colonization from thesouth as well as from the east. The patterns found in the American populationsare consistent with post-glacial colonization, possibly even with anthropogenicintroduction from Europe. Our data also suggest that the tussock-formingpopulations of C. nigra, often referred to as a distinct species (Carex juncella),represent an ecotype that has originated repeatedly from different populationswith creeping rhizomes.KeywordsBoreal–temperate herb, Europe, extensive gene flow, glacial refugia, rpl32–trnL