Plant dependence on fungal carbon (mycoheterotrophy) evolved repeatedly. In orchids, it is connected with a mycorrhizal shift from rhizoctonia to ectomycorrhizal fungi and a high natural (13)C and (15)N abundance. Some green relatives of mycoheterotrophic species show identical trends, but most of these remain unstudied, blurring our understanding of evolution to mycoheterotrophy. We analysed mycorrhizal associations and (13)C and (15)N biomass content in two green species, Neottia ovata and N. cordata (tribe Neottieae), from a genus comprising green and nongreen (mycoheterotrophic) species. Our study covered 41 European sites, including different meadow and forest habitats and orchid developmental stages. Fungal ITS barcoding and electron microscopy showed that both Neottia species associated mainly with nonectomycorrhizal Sebacinales Clade B, a group of rhizoctonia symbionts of green orchids, regardless of the habitat or growth stage. Few additional rhizoctonias from Ceratobasidiaceae and Tulasnellaceae, and ectomycorrhizal fungi were detected. Isotope abundances did not detect carbon gain from the ectomycorrhizal fungi, suggesting a usual nutrition of rhizoctonia-associated green orchids. Considering associations of related partially or fully mycoheterotrophic species such as Neottia camtschatea or N. nidus-avis with ectomycorrhizal Sebacinales Clade A, we propose that the genus Neottia displays a mycorrhizal preference for Sebacinales and that the association with nonectomycorrhizal Sebacinales Clade B is likely ancestral. Such a change in preference for mycorrhizal associates differing in ecology within the same fungal taxon is rare among orchids. Moreover, the existence of rhizoctonia-associated Neottia spp. challenges the shift to ectomycorrhizal fungi as an ancestral pre-adaptation to mycoheterotrophy in the whole Neottieae.

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Summary Arbuscular mycorrhizal fungi (AMFs) are important plant symbionts, but we know little about the effects of plant taxonomic identity or functional group on the AMF community composition. To examine the effects of the surrounding plant community, of the host, and of the AMF pool on the AMF community in plant roots, we manipulated plant community composition in a long‐term field experiment. Within four types of manipulated grassland plots, seedlings of eight grassland plant species were planted for 12 wk, and AMFs in their roots were quantified. Additionally, we characterized the AMF community of individual plots (as their AMF pool) and quantified plot abiotic conditions. The largest determinant of AMF community composition was the pool of available AMFs, varying at metre scale due to changing soil conditions. The second strongest predictor was the host functional group. The differences between grasses and dicotyledonous forbs in AMF community variation and diversity were much larger than the differences among species within those groups. High cover of forbs in the surrounding plant community had a strong positive effect on AMF colonization intensity in grass hosts. Using a manipulative field experiment enabled us to demonstrate direct causal effects of plant host and surrounding vegetation.

Arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSE) form symbiotic relationships with plants influencing their productivity, diversity and ecosystem functions. Only a few studies on these fungi, however, have been conducted in extreme elevations and none over 5500 m a.s.l., although vascular plants occur up to 6150 m a.s.l. in the Himalayas. We quantified AMF and DSE in roots of 62 plant species from contrasting habitats along an elevational gradient (3400–6150 m) in the Himalayas using a combination of optical microscopy and next generation sequencing. We linked AMF and DSE communities with host plant evolutionary history, ecological preferences (elevation and habitat type) and functional traits. We detected AMF in elevations up to 5800 m, indicating it is more constrained by extreme conditions than the host plants, which ascend up to 6150 m. In contrast, DSE were found across the entire gradient up to 6150 m. AMF diversity was unimodally related to elevation and positively related to the intensity of AMF colonization. Mid-elevation steppe and alpine plants hosted more diverse AMF communities than plants from deserts and the subnival zone. Our results bring novel insights to the abiotic and biotic filters structuring AMF and DSE communities in the Himalayas.

For the first time in bryophyte studies, we performed comprehensive cloning of the ITS region to reveal intraindividual variation of ITS sequences. We assessed relationships among morphologically defined taxa of the polyploid complex of the moss Tortula muralis. Our results detected a monophyletic T. muralis complex comprising T. muralis subsp. mura lis, T. muralis subsp. obtusifolia, T. lingulata, T. israelis, and T. edentula. The single accession of T. edentula was found nested within T. obtusifolia, and biphyletic T. israelis was found to be nested within T. muralis. With the exception of T. lingulata, intragenomic ITS sequence variation was high in the T. muralis complex. Most intraindividual sequences were nevertheless only weakly divergent, suggesting their origin via mutations exceeding the rates of concerted evolution. Markedly divergent sequences found within a single individual most probably resulted from gene flow among distant lineages of the complex. Such pattern of ITS variation challenges the traditional morphology-based taxonomy. No phylogenetic signal was associated with ploidy-level variation, suggesting a polytopic origin of the diploids. Interestingly, the pattern of ITS variation together with morphological evidence indicate the autopolyploid origin of some lineages, which renders the T. muralis complex the first group of mosses in which autopolyploidy is implied by molecular markers.

Abstract:The ability to grow clonally is generally considered important for plants in Arctic regions but analyses of clonal characteristics are lacking for entire plant communities. To fill this gap, we assessed the clonal growth of 78 plant species in the Petuniabukta region, central Spitsbergen (Svalbard), and analyzed the clonal and other life−history traits in the re− gional flora and plant communities with respect to environmental gradients. We distin− guished five categories of clonal growth organs: perennial main roots produced by non− clonal plants, epigeogenous rhizomes, hypogeogenous rhizomes, bulbils, and stolons. Clonal growth differed among communities of the Petuniabukta region: non−clonal plants prevailed in open, early−successional communities, but clonal plants prevailed in wetlands. While the occurrence of plants with epigeogenous rhizomes was unrelated to stoniness or slope, the occurrence of plants with hypogeogenous rhizomes diminished with increasing stoniness of the substratum. Although the overall proportion of clonal plants in the flora of the Petuniabukta region was comparable to that of central Europe, the flora of the Petunia− bukta region had fewer types of clonal growth organs, a slower rate of lateral spread, and a different proportion of the two types of rhizomes.

Ranunculus section Batrachium (water crowfoot) ranks among the most taxonomically challenging aquatic plant groups due to morphological reduction, phenotypic plasticity, polyploidy and reticulate evolution. This study, for the first time in this group, linked morphology, genome size and genetic data (two non-coding regions of plastid DNA and the ITS region of nuclear ribosomal DNA). This extensive data set, including 258 central European populations, enables us to recognize widespread lineages from hybrids and to trace the evolutionary processes underlying the observed diversity. Most of the traditionally recognized species are supported. However, the presence of two morphologically cryptic but genetically well-differentiated lineages was detected within Ranunculus trichophyllus, and three separate lineages of different parentage were confirmed in the Ranunculus penicillatus complex. An allopolyploid origin was revealed in Ranunculus aquatilis, Ranunculus baudotii and in both lineages of R. trichophyllus, for which the parentage has not yet been studied, and allopolyploidy is suspected in all other polyploid taxa except for the triploid cytotype of Ranunculus fluitans, which is most likely autopolyploid. We detected putative F1 hybrids of seven different parentage combinations, including two involving Ranunculus rionii, representing the first known hybrids of this species. An additional 13 hybrid cytotypes (backcrosses or entailing additional polyploidization) were revealed; c. 20% of Ranunculus peltatus accessions seem to be influenced by introgression detectable only using sequence data. The Batrachium group is remarkable due to the coexistence of well-defined lineages with recently evolved biotypes arising due to hybridization and polyploidization.

Blastenia is a widely distributed lichen genus in Teloschistaceae. We reconstructed its phylogeny in order to test species delimitation and to find evolutionary drivers forming recent Blastenia diversity. The origin of Blastenia is dated to the early Tertiary period, but later diversification events are distinctly younger. We recognized 24 species (plus 2 subspecies) within 6 infrageneric groups. Each species strongly prefers a single type of substrate (17 species occur on organic substrates, 7 on siliceous rock), and most infrageneric groups also show a clear substrate preference. All infrageneric groups tend to have the Mediterranean and Macaronesian distribution, but some epiphytic species have much larger geographic ranges and some evolved after a long-distance dispersal outside the region. Chlorinated and nonchlorinated anthraquinone chemosyndromes co-occur in apothecia of most species, but the chemotype has been secondarily reduced in some lineages. One infrageneric group has a marked reduction in apothecial size, associated with a substrate shift to twigs. Only seven species have vegetative diaspores; they also produce apothecia but have smaller ascospores. Genome sizes (22-35 Mb in Blastenia) are significantly higher in epilithic species. Within-species genetic variation is low in widely distributed species but high in some epilithic species with small geographical ranges. New taxa are: B. afroalpina, B. anatolica, B. caucasica, B. gennargentuae, B. herbidella subsp. acidophila, B. lauri, B. monticola, B. palmae, B. psychrophila, B. purpurea, B. relicta, B. remota, B. xerothermica, and B. xerothermica subsp. macaronesica. New combinations are: B. festivella and B. subathallina; both names and B. catalinae are lectotypified.