Tiina Randlane

Evolution of complex symbiotic relationships in a morphologically derived family of lichen-forming fungi.

We studied the evolutionary history of the Parmeliaceae (Lecanoromycetes, Ascomycota), one of the largest families of lichen-forming fungi with complex and variable morphologies, also including several lichenicolous fungi. We assembled a six-locus data set including nuclear, mitochondrial and low-copy protein-coding genes from 293 operational taxonomic units (OTUs). The lichenicolous lifestyle originated independently three times in lichenized ancestors within Parmeliaceae, and a new generic name is introduced for one of these fungi. In all cases, the independent origins occurred c. 24 million yr ago. Further, we show that the Paleocene, Eocene and Oligocene were key periods when diversification of major lineages within Parmeliaceae occurred, with subsequent radiations occurring primarily during the Oligocene and Miocene. Our phylogenetic hypothesis supports the independent origin of lichenicolous fungi associated with climatic shifts at the Oligocene-Miocene boundary. Moreover, diversification bursts at different times may be crucial factors driving the diversification of Parmeliaceae. Additionally, our study provides novel insight into evolutionary relationships in this large and diverse family of lichen-forming ascomycetes.

World survey of the genus Lepraria ( Stereocaulaceae , lichenized Ascomycota)

A comparative review of 57Lepraria species and 2 varieties is provided together with species descriptions and a key. Lecanora leuckertiana is transferred to Lepraria. In addition some putative taxa by different authors are discussed. ducing anthraquinones to Caloplaca Th. Fr. and Leproplaca (Nyl.) Nyl. ex Hue (a current synonym of Caloplaca) and species contain- ing pulvinic acid derivatives to Chrysothrix Mont. The number of species in Lepraria was further reduced when taxa producing dibenzofuranes were placed in Leproloma Nyl. ex Cromb. (Laundon 1989; Leuckert & Kummerling 1991). The concept of the

Phylogeny of the cetrarioid core (Parmeliaceae) based on five genetic markers

Fourteen genera belong to a monophyletic core of cetrarioid lichens, Ahtiana, Allocetraria, Arctocetraria, Cetraria, Cetrariella, Cetreliopsis, Flavocetraria, Kaernefeltia, Masonhalea, Nephromopsis, Tuckermanella, Tuckermannopsis, Usnocetraria and Vulpicida. A total of 71 samples representing 65 species (of 90 worldwide) and all type species of the genera are included in phylogentic analyses based on a complete ITS matrix and incomplete sets of group I intron, β-tubulin, GAPDH and mtSSU sequences. Eleven of the species included in the study are analysed phylogenetically for the first time, and of the 178 sequences, 67 are newly constructed. Two phylogenetic trees, one based solely on the complete ITS-matrix and a second based on total information, are similar, but not entirely identical. About half of the species are gathered in a strongly supported clade composed of the genera Allocetraria, Cetraria s. str., Cetrariella and Vulpicida. Arctocetraria, Cetreliopsis, Kaernefeltia and Tuckermanella are monophyletic genera, whereas Cetraria, Flavocetraria and Tuckermannopsis are polyphyletic. The taxonomy in current use is compared with the phylogenetic results, and future, probable or potential adjustments to the phylogeny are discussed. The single non-DNA character with a strong correlation to phylogeny based on DNA-sequences is conidial shape. The secondary chemistry of the poorly known species Cetraria annae is analyzed for the first time; the cortex contains usnic acid and atranorin, whereas isonephrosterinic, nephrosterinic, lichesterinic, protolichesterinic and squamatic acids occur in the medulla. Notes on the anatomy of Cetraria annae and Flavocetraria minuscula are also provided.

The cetrarioid core group revisited (Lecanorales: Parmeliaceae)

The cetrarioid core group has been the focus of numerous taxonomic and phylogenetic studies in recent years, yet the phylogenetic resolution and support among these clades remains unclear. Here we use four commonly employed loci to estimate if their use increases phylogenetic resolution and support. The present study largely confirms the topologies of previous studies, but with increased support. Approximately half of the genera in the cetrarioid core were not monophyletic. Melanelia sorediella was clustered within Cetrariella, and the combination Cetrariella sorediella (Lettau) V. J. Rico & A. Thell comb. nov. is made. Additionally, the genus Flavocetrariella was supported as part of Nephromopsis and is considered to be a synonym of the latter. Finally, a comparison of genetic distances shows that the maximum intrageneric genetic distance encompassed by many cetrarioid genera is lower than that of many other genera in Parmeliaceae.

A second, updated world list of cetrarioid lichens

At the present time 135 cetrarioid lichen species can be listed. According to recent revisionary work they are placed into 22 genera. Six new genera (Arctocetraria, Cetrariella, Flavocetraria, Kaernefeltia, Nimisia and Tuckneraria) have been separated since 1993 when the first world list of cetrarioid lichens was presented. The genus Cetrariopsis is included in Nephromopsis in this paper. An alphabetical list of all epithets that have been applied to cetrarioid lichens is given, and the generic name to which they have been assigned is indicated in brackets. The author(s) of each combination, as well as the issue of its valid publication, together with the page and year are also presented. The currently accepted combination is indicated for each epithet. Two new combinations are proposed: Nephromopsis pallescens var. citrina (Taylor) Thell & Randlane and Nephromopsis laii (Thell & Randlane) Saag & Thell. (Less)

The lichen genus Usnea (lichenized Ascomycetes, Parmeliaceae) in Estonia with a key to the species in the Baltic countries

The occurrence of 12 Usnea species in Estonia is verified by investigating c . 1800 specimens. The known distribution and substratum preferences of Usnea taxa in Estonia are summarized. Picea is the main substratum for Usnea species in Estonia, but occasionally these species occur also on deciduous trees, wood, and exceptionally on rock. A key for identification of 16 taxa recorded in the three Baltic countries is provided and diagnostic morphological and chemical characters used in the key are discussed. A new chemotype of U. fulvoreagens has been identified. U. wasmuthii is reported as new to Lithuania.

Phylogenetic relations of European shrubby taxa of the genus Usnea

This study focuses on European Usnea species with sorediate shrubby thalli, with the aim to evaluate the morphological and chemical separation of species in the light of molecular data. Twenty-two Usnea species, including widely distributed taxa such as U. diplotypus, U. fulvoreagens, U. glabrescens, U. lapponica, U. subfloridana, U. substerilis and U. wasmuthii, were included in the study using Bayesian and maximum parsimony analyses of nuclear ITS and beta-tubulin sequences. The analyses showed that: 1) most taxa that are morphologically well delimited are also distinct by means of molecular characters, 2) shrubby taxa in the section Usnea that are difficult to determine by traditional characters form a group of closely related but still genetically distinct entities, except U. diplotypus and U. substerilis which appear to be polyphyletic. The branch lengths differed largely between two parts of the ITS tree (sections Usnea and Ceratinae). Usnea intermedia is proposed as the sexually reproducing counterpart for the sorediate U. lapponica. Additionally, some new chemotypes of Usnea species were determined.

A new circumscription of the lichen genus Nephromopsis (Parmeliaceae, lichenized Ascomycetes).

The phylogeny of the cetrarioid lichens with bifusiform spermatia and dorsiventral thalli which contain usnic acid is reanalysed using three parts of the genome, ITS rDNA, β-tubulin and GAPDH sequences. Molecular data from five cetrarioid species are presented for the first time, and 13 new sequences are combined with sequences from the gene bank to delimit the genus Nephromopsis. A monophyletic clade of Nephromopsis, Tuckneraria, ‘Cetraria’ leucostigma and ‘C.’ melaloma is identified and circumscribed as one genus, Nephromopsis, which now includes 19 species. Four new combinations are presented. A key to the species is provided.

Species delimitation in the lichenized fungal genus Vulpicida (Parmeliaceae, Ascomycota) using gene concatenation and coalescent-based species tree approaches.

UNLABELLED • PREMISE OF THE STUDY Species boundaries in many organism groups are still in a state of flux, and for empirical species delimitation, finding appropriate character sets and analytical tools are among the greatest challenges. In the lichenized fungal genus Vulpicida, six morphologically circumscribed species have been distinguished, but phenotypic characters partly overlap for three of these and intermediate forms occur. We used a combination of phylogenetic strategies to delimit the species in this genus.• METHODS Five DNA loci were sequenced and analyzed. Single-locus gene trees and a five-locus concatenated phylogeny were constructed to assess current Vulpicida species. Species boundaries were inferred from molecular data using two coalescent-based species delimitation methods (BP&P and Brownie) and from species trees reconstructed with three different algorithms (*BEAST, BEST, and STEM).• KEY RESULTS The two species restricted to North America, Vulpicida canadensis and V. viridis, are clearly distinct in all analyses. The four other traditionally accepted species form two strongly supported, closely related species-level lineages within the core group of the genus. On the basis of these results, we propose four instead of the current six species in the genus: V. canadensis, V. juniperinus, V. pinastri, and V. viridis, while V. tilesii and V. tubulosus are reduced to synonymy under V. juniperinus.• CONCLUSIONS Coalescent species delimitation and tree inference give consistent results for fully distinct Vulpicida species but not for diverging populations. Even the inconsistent results were informative, revealing developing isolation despite a complex history of recombination and incomplete lineage sorting.

Erratum to: Evaluation of traditionally circumscribed species in the lichen-forming genus Usnea , section Usnea (Parmeliaceae, Ascomycota) using a six-locus dataset

The MycoBank code of the newly described species Usnea parafloridana was missing. The MycoBank code for the species Usnea parafloridana should read: MB818694. Usnea parafloridana K. Mark, Will-Wolf & Randlane sp. nov.; MycoBank No. MB818694. Type: USA, Wisconsin, Vilas Co., Trout Lake Conifer Swamp State Natural Area; 46.0135° N, -89.6586° W; 27.08.2011, Susan Will-Wolf WW14807: isolates WW_018 (holotype, TU; Fig. 4a, c, e), WW_023 (isotype 1, WIS), WW_013 (isotype 2, F). Morphology: thallus shrubby, up to 3–6 cm long, often with relatively few branches; branching mainly isotomic-dichotomous, divergent; lateral branches not narrowed at point of attachment; basal part distinctly jet black, with few annular cracks; papillae verrucose, numerous on main branches and lesser or absent on lateral branches; fibrils few to numerous; soralia small and punctiform when young, enlarging, becoming close to each other but usually staying delimited when mature, more numerous on terminal branches; isidiomorphs numerous, spinulose, relatively short and thick, both on young and mature soralia; cortex thick (9–15%); medulla thin (10.5– 13%), dense, not pigmented; central cord thick (60–73%) and white; apothecia not seen (Fig. 4; colour illustrations in online version). Secondary chemistry: usnic acid in cortex; norstictic acid as a major compound, salazinic acid as an accessory substance (present in most examined specimens) in medulla. Ecology: on branches of Abies balsamea, Larix laricina, Picea mariana, or Pinus strobus in cedar swamp, conifer bog and pine plantation with trees over one-hundred years old. Distribution: currently 15 specimens are known from four localities in Wisconsin, USA. Etymology: the species is morphologically somewhat similar to Usnea subfloridana (both taxa have similar shrubby thalli, black basal parts and delimited soralia with numerous isidiomorphs), which phylogenetically appears conspecific with U. florida. The same root ‘florida’ is used in the epithet of the new taxon to underline this morphological similarity while the prefix ‘para-‘ indicates phylogenetic distinctness of the species from U. florida and U. subfloridana. The online version of the original article can be found at http://dx.doi. org/10.1007/s13127-016-0273-7.