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We tested whether the conspicuous lichen Lobaria pulmonaria indicates the number of tree-dependent, red-listed species in a hemiboreal forest in southern Sweden. In 18 naturally regenerated, mainly old deciduous forest plots considered to be of high or very high conservation value, the number of red-listed tree lichens or wood beetles was not positively correlated with the area of the forest stands studied (8–56 ha). The 8 stands with L. pulmonaria had about nine (median) red-listed lichens, but 10 stands without L. pulmonaria had only about one such species, a highly significant difference. Similarly, the variation between stands in the number of red-listed wood beetles was considerable, but it was very weakly correlated with the number of red-listed lichens. The number of red-listed wood beetles dependent on dead trees was not different in areas with or without Lobaria pulmonaria, but the number of red-listed wood beetles dependent on hollow trees was higher in stands with Lobaria pulmonaria (median of seven species) than in those without (three species) the lichen. Stands with this lichen species also contained significantly more of other types of lichens that have been proposed as indicators of forest continuity according to three lists. We stress the need for identification of species that could serve as indicators of different types of forest continuity and identify some organism groups that may indicate the different types. Los liquenes y coleopteros dependientes de arboles como indicadores en los bosques bajo conservacion Resumen: Evaluamos si el consipicuo liquen Lobaria pulmonaria indica el numero de especies dependientes de los arboles, listadas en el libro rojo en un area de bosque hemi-boreal en el sur de Suecia. El numero de liquenes de los arboles o coleopteros de la madera listados en el libro rojo no estuvo positivamente correlacionado con el area de los rodales de bosque estudiados (8–56 ha), en 18 rodales naturalemente regenerados, en su mayoria de antiguo crecimiento de bosques deciduos; los cuales se considera tienen de alto a muy alto valor de conservacion. Los 8 rodales con L. pulmonaria tuvieron alrededor de nueve (mediana) liquenes listados en el libro rojo, pero 10 rodales sin L. pulmonaria tuvieron solamente una de dichas especies, una diferencia significativamente alta. Asi mismo, la variacion entre rodales en el numero de coleopteros listados en el libro rojo fue considerable, pero estuvo debilmente correlacionada con el numero de liquenes listados en el libro rojo. El numero de coleopteros de la madera listados en el libro rojo, dependientes de arboles muertos no fue diferente en areas con o sin Lobaria pulmonaria. Sin embargo, el numero de coleopteros de la madera listados en el libro rojo dependientes de troncos huecos fue mas alto en rodales con Lobaria pulmonaria (mediana = siete especies) que en aquellos sin el mencionado liquen (tres epecies). Los rodales con este liquen tambien contuvieron significativamente un numero mayor de otros liquenes que han sido propuestos como indicadores de la continuidad del bosque de acuerdo a tres listas. Remarcamos la necesidad de la identificacion de especies que pueden servir como indicadoras de los diferentes tipos de continuidad del bosque y la identificacion de ciertos grupos de organismos que pueden indicar los diferentes tipos.

Species identification in the tropical lichen genusPhyllopsorais generally challenging and is based on ascospore morphology, vegetative dispersal units, thallus structure and secondary chemistry. As several type specimens are in poor condition and difficult to interpret, it is often unclear how these old names fit with the currently used taxonomy. In the present study, we aim to identify species boundaries inPhyllopsoras. str. supported by an integrative approach using multiple sources of evidence. We investigated a substantial amount of herbarium as well as freshly collected material and generated mtSSU and ITS sequence data from most of the described species, including several types. Species delimitation analyses are applied on the gene trees using mPTP and we construct a species tree of both markers with *BEAST, facilitating discussion of species delimitation and sister-relationships. Comparing morphology, chemistry and molecular data, we found that the mPTP analyses split established species repeatedly. Based on our integrative results, we exclude nine species from the genus, resurrect one (P. melanoglaucaZahlbr.), reduce two into synonymy with otherPhyllopsoraspecies and describe five as new to science:Phyllopsora amazonicaKistenich & Timdal (which shares the secondary chemistry (atranorin and terpenoid pattern) withP. haleichemotype 1, but differs, e.g., in having smaller areolae that are attached to a thinner, white prothallus, and in having more persistently marginate and less convex apothecia),Phyllopsora concinnaKistenich & Timdal (which shares the secondary chemistry (atranorin and parvifoliellin) withP. parvifoliellaandP. rappiana, but differs from both in forming larger isidia, having a white prothallus, apothecial margin paler than the disc, and longer and broader ascospores),Phyllopsora furfurellaKistenich & Timdal (which is here segregated fromP. furfuraceabased on having a white prothallus and in containing skyrin in the hypothecium (K+ red)),Phyllopsora isidosaKistenich & Timdal (which differs fromP. byssisedain forming a more crustose thallus with more delicate isidia, and fromP. isidiotylain forming somewhat coarser, less branched isidia) andPhyllopsora neotinicaKistenich & Timdal (a neotropical species here segregated from the now exclusively paleotropicalP. chodatinica, differing in containing an unknown xanthone (not chodatin)). Lectotypes are designated forBiatora pyrrhomelaenaTuck.,Lecidea leucophyllinaNyl.,L. pertextaNyl., andP. brachysporaMüll. Arg. In total, we accept 54 species in the genusPhyllopsora.

BiatoraalnetorumS. Ekman & Tønsberg, a lichenised ascomycete in the family Ramalinaceae (Lecanorales, Lecanoromycetes), is described as new to science. It is distinct from other species ofBiatorain the combination of mainly three-septate ascospores, a crustose thallus forming distinctly delimited soralia that develop by disintegration of convex pustules and the production of atranorin in the thallus and apothecia. The species is known from the Pacific Northwest of North America, where it inhabits the smooth bark of Alnusalnobetulasubsp.sinuata andA.rubra.Biatoraalnetorumis also a new host for the lichenicolous ascomyceteSclerococcumtoensbergiiDiederich.

Fellhaneropsis almquistiorum S. Ekman, a lichenized ascomycete in the family Pilocarpaceae (Lecanorales, Lecanoromycetes) is described as new to science. It is characterized by the thin and filmy crustose thallus, small photobiont, apothecia with a yellowish to black disk and well developed black margin that becomes excluded with age, dark green and brown pigmentation in the proper exciple and hypothecium, a hymenium without pigment or with some green and brown pigment, more or less fusiform and 3-septate ascospores, abundant black pycnidia, and long and filiform conidia. The species is currently known from central Sweden and central Germany, where it inhabits shaded and acid to neutral rocks with few competing lichens. Historical records were all misidentified as Bacidia coprodes or any of its synonyms or misapplied names. Some specimens of F. almquistiorum contain a previously unrecorded pigment, tentatively named here Almquistiorum-yellow. This pigment is intensely golden yellow in water, does not react with KOH and fades with the addition of HCl or HNO3. A key to all known, pycnidiate members of Fellhaneropsis is provided.

The Ramalinaceae is the fourth‐largest family of lichenized ascomycetes with 42 genera and 913 species exhibiting considerable morphological variation. Historically, generic boundaries in the Ramalinaceae were primarily based on morphological characters. However, molecular systematic investigations of subgroups revealed that current taxonomy is at odds with evolutionary relationships. Tropical members of the family remain particularly understudied, including the large genus Phyllopsora. We have generated and collected multilocus sequence data (mtSSU, nrITS, nrLSU, RPB1, RPB2) for 149 species associated with the Ramalinaceae and present the first comprehensive molecular phylogeny of the family. We used ancestral state reconstructions on our molecular family phylogeny to trace the evolution of character states. Our results indicate that the Ramalinaceae have arisen from an ancestor with long, multiseptate ascospores living in humid temperate forests, and that the phyllopsoroid growth form has evolved multiple times within the family. Based on our results using integrative taxonomy, we discuss sister‐relations and taxon‐delimitation within five well‐supported clades: The Bacidia, Biatora‐, Ramalina‐, Rolfidium‐, and Toninia‐groups. We reduce six genera into synonymy and make 49 new nomenclatural combinations. The genera Bacidia, Phyllopsora, Physcidia and Toninia are polyphyletic and herein split into segregates. We describe the two genera Bellicidia and Parallopsora and resurrect the genera Bibbya, Kiliassia, Sporacestra, and Thalloidima. According to our new circumscription, which also includes some additional changes, the family Ramalinaceae now comprises 39 genera. Locked until 18.8.2019 due to copyright restrictions. This is the peer reviewed version of an article, which has been published in final form at [https://doi.org/10.12705/675.1]. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving.

We present taxonomic, distributional and ecological notes on Fennoscandian crustose lichens and lichenicolous fungi, based on new collections as well as revision of herbarium material. Two new combinations are proposed: Frutidella furfuracea comb. nov. for F. pullata and Puttea duplex comb. nov. for Fellhanera duplex. Lecidea byssoboliza, L. carneoglauca and Variolaria torta are all reduced to synonymy with Bacidia antricola, Bacidia invertens is synonymized with B. igniarii, B. atrolivida with Mycobilimbia tetramera, and Gyalidea fruticola with Thelenella pertusariella. A new description is provided for Micarea hylocomii. 25 species of lichens and lichenicolous fungi are reported as new to Finland, Norway and/or Sweden: Absconditella lignicola (Norway), Bacidia antricola (Norway), B. polychroa (Norway), B. pycnidiata (Sweden), Bacidina adastra (Sweden), Biatora veteranorum (Norway), Briancoppinsia cytospora (Finland), Catillaria scotinodes (Norway), Cliostomum subtenerum (Norway), Dirina fallax (Sweden), Fellhaneropsis almquistiorum (Norway), Gyalidea subscutellaris (Sweden), Lecania inundata (Norway), L. suavis (Norway), Micarea capitata (Norway), M. deminuta (Norway), M. hylocomii (Sweden), M. lynceola (Sweden), M. soralifera (Sweden), M. subconfusa (Sweden), Mycoblastus sanguinarioides (Finland, Sweden), Paralecia pratorum (Sweden), Puttea duplex (Sweden), Sarcogyne algoviae (Finland) and Toninia subnitida (Norway). Lectotypes are designated for Bacidia antricola, Lecidea byssoboliza, Lecidea carneoglauca, Lecidea subconfusa and Lecidea submoestula.

We aim to increase understanding and improve taxonomy of seven names currently treated in Bacidia that were described from Russian territory based on material collected during 19th-century expeditions: Bacidia alborussula , B. freshfieldii , B. graminum , B. indigens , B. primigenia , B. subabbrevians , and B. xylophila . B. alborussula , B. graminum , and B. indigens are transferred here to Haematomma , Lecania , and Bacidina , respectively. Bacidia freshfieldii and B. subabbrevians belong in the Ramalinaceae and are provisionally kept in Bacidia even though none of them is congeneric with the type of that genus. B. primigenia belongs in Arthrorhaphis , but we refrain from making any new combination owing to its questionable status as a species distinct from A. grisea . B. xylophila is a younger synonym of Lecania subfuscula . We designate lectotypes for names previously not typified and for which syntypes exist: Lecidea alborussula , L. graminum , L. indigens , and L. subabbrevians .

The new generic name Bryobilimbia is proposed for Lecidea hypnorum and the closely related taxa Lecidea ahlesii var. ahlesii, L. ahlesii var. nemoralis, L. diapensiae, L. sanguineoatra and Mycobilimbia australis. A phylogenetic analysis based on five genes shows that Lecidea berengeriana does not belong to this group but is more closely related to Romjularia. Both groups of species have been included in Mycobilimbia by some authors but are instead shown to be most closely related to a group of genera (including Clauzadea, Farnoldia, Lecidoma and Romjularia) that do not belong to Lecideaceae s. str. A neotype is selected for Lichen sanguineoater Wulfen and the new combinations Bryobilimbia ahlesii var. ahlesii, B. ahlesii var. nemoralis, B. australis, B. diapensiae, B. hypnorum and B. sanguineoatra are proposed. A lectotype is also selected for L. templetonii Taylor.

A taxonomic and biogeographic overview of the genusMyrionorais provided. Two species are recognized,M. albidula(Willey) R. C. Harris andM. pseudocyphellariae(Etayo) S. Ekman & Palice comb. nov. The genus is characterized by polysporous asci, the presence of crystals in the hymenium and proper exciple that partly consist of lobaric acid, and a photobiont with large cells (mostly in the range 12–20 µm).Myrionora albidulais currently known from Germany, Norway, Sweden, the Russian Federation (Altayskiy Kray, Chelyabinskaya Oblast', Khabarovskiy Kray and Zabaykal'skiy Kray), and the United States (Alaska, Connecticut, Maine and Massachusetts). It inhabits bark of deciduous trees and shrubs and conifers over a wide range of latitudes.Myrionora pseudocyphellariaeis known from Chile and Ecuador, where it has been encountered on lichens and decaying bark. Based on morphological characteristics, we conclude thatMyrionorabelongs in theRamalinaceae.

North European epiphytic lichens are often genetically impoverished compared with their North American counter- parts. This has been hypothesized to impede sexual reproduction due to reduced chances of finding compatible mating type partners. We compared genetic variation and reproductive mode in two threatened Scandinavian lichens, Evernia divaricata and Usnea longissima, with more viable populations in North America to see (i) if these species also show genetical depletion in northern Europe and (ii) if the occurrence of sexual propagules (ascospores in apothecia) is more prevalent in genetically diverse populations. Genetic variation of the fungal component was assessed by sequencing two nuclear rDNA gene regions (ITS and IGS) in 1005 and 1477 thalli, collected from 92 and 160 localities of E. divaricata and U. longissima, respectively. Scandinavian popula- tions of both species were almost devoid of genetic variation compared with much higher genetic diversity in North America. We found no support for the proposed relationship between genetic diversity and fertility. Fertile thalli were found in several genetically invariable populations. Fertility increased with population size and regional abundance in E. divaricata, but not in U. longissima. In Scandinavia, E. divaricata was more fertile than previously recorded, whereas all sampled populations of U. longissima were sterile and possibly clonal. Resume : Les lichens epiphytes europeens sont souvent appauvris genetiquement, comparativement aleurs contreparties americaines. On a formule l'hypothese que ceci nuirait ala reproduction sexuelle, du ala reduction des chances de trouver un partenaire de type de croisement compatible. Les auteurs ont compare la variation genetique et le mode reproducteur chez deux lichens scandinaves menaces, l'Evernia divaricata et l'Usnea longissima, avec des populations plus viables de l'Amerique du Nord afin de verifier (i) si ces especes montrent egalement un epuisement genetique dans le nord de l'Europe et (ii) si la presence de propagules sexuelles (ascospores dans des apotheces) prevaut plus fortement dans des populations genetiquement diverses. Les auteurs ont evalue la variation genetique de la composante fongique en sequencant deux regions du gene nucleique tADN (ITS et IGS) chez 1005 et 1477 thalles recoltes a partir de 92 et 160 localites, pour l'E. divaricata et l'U. longissima, respectivement. Les populations scandinaves des deux especes sont pratiquement depourvues de variation genetique comparativement aune diversite genetique tres fortement marquee en Amerique du Nord. Il n'ont trouve aucun support pour la proposition suggerant une relation entre la diversite genetique et la fertilite. On a retrouve des thalles fertiles dans plusieurs populations genetique- ment invariables. La fertilite augmente avec la dimension de la population chez l'E. divaricate, mais non chez l'U. longisima .E n Scandinavie, l'E. divaricata s'avere plus fertile que reconnu jusqu'ici, alors que toute les populations echantillonnees de l'U. longissima se sont averees steriles et possiblement clonales. (Traduit par la Redaction) Mots-cles : richesse allelique, dispersion, lichens epiphytes, diversite genetique, forets surannees.

New records, range extensions and nomenclatural innovations for lichens and lichenicolous fungi from Alaska, U.S.A. - Herzogia 25: 177-210. Surveys of lichens and lichenicolous fungi have been taki ...

Morphological characters can vary to an extent that makes it difficult to separate the two morphologically and chemically similar species Xanthoria aureola and X. parietina for an inexperienced field biologist. They occur frequently in mixed stands on seashore rocks on the western coast of Norway. We examined (1) whether a simple method like RAPD-PCR could confirm the distinction between the two species previously reported on the basis of DNA sequences, and (2) whether infraspecific DNA and RAPD data from X. parietina were congruent. We also checked whether the RAPD band scoring procedure could affect the results. Results show that (1) RAPD distance matrices based on band scorings performed independently by the two authors were always congruent and differences never affected conclusions, (2) RAPD clearly distinguishes between X. parietina and X. aureola in a way that is fully congruent with a classification based on DNA sequence data, and (3) there was no significant congruence between infraspecific distances based on DNA sequences and RAPD data in X. parietina. The latter observation may be taken as support for a previously published report of low levels of recombination in X. parietina, which stands in contrast to statements of obligate homothallism in that species.

The recently described genus Vahliella (Peltigerales, Ascomycetes) has repeatedly appeared outside the Pannariaceae in molecular phylogenies. Here we include data from additional species of the genus and utilize mtSSU rDNA and RPB1 sequences to confirm its placement as the sister to a group consisting of Lobariaceae, Massalongiaceae, Nephromataceae and Peltigeraceae, in the Peltigerales. The new family Vahliellaceae Wedin, P. M. Jørg. & S. Ekman is described for the genus, and its morphological characteristics are briefly discussed.

Recent surveys of the inland rain forests of British Columbia and adjacentregions have brought to light an unexpectedly rich epiphytic lichen flora, including severalspecies apparently new to scien ...

In order to examine genetic variation and population structure of the widespread lichen-forming ascomycete Xanthoria parietina from similar habitats, but different sites in Scandinavia, we investigated seven populations in Scania, southernmost Sweden, and compared the results with a corresponding study on Storfosna, central Norway. Sequence variations of the nuclear ribosomal DNA were used as molecular markers, for both a part of the IGS region and the complete ITS1-5.8S-ITS2 region. The amount of genetic variability observed was comparable in the two investigations. Divergence between populations in different habitats found in the previous study was also present in this study. Xanthoria parietina is genetically differentiated between habitats with no evidence of restricted gene flow between populations in the same habitat at the present spatial scale, at least at sites along the coast of Scandinavia. Differentiation between habitats is considerable at both study sites, which we attribute to restricted gene flow between habitats, i.e. habitat isolation.

The family Parmeliaceae (Lecanorales, Ascomycota) is possibly the largest, best known and most thoroughly studied lichen family within its order. Despite this fact the relationship between Parmeliaceae and other groups in Lecanorales is still poorly known. The aim of the present study is to contribute to finding the sister group of Parmeliaceae as an aid in future studies on the phylogeny and character evolution of the group. We do this by sampling all potential relatives to the Parmeliaceae that we have identified, i.e. Gypsoplaca, Japewia, Mycoblastus, Protoparmelia, and Tephromela, a good representation of the major groups within the Parmeliaceae s. lat. and a good representation of other taxa in the core Lecanorales. We use molecular data from two genes, the large subunit of the nuclear ribosomal RNA gene (nrLSU) and the small subunit of the mitochondrial ribosomal RNA gene (mrSSU), and a Bayesian analysis of the combined data. The results show that the closest relatives to Parmeliaceae are the two genera Protoparmelia and Gypsoplaca, which are crustose lichens. Parmeliaceae in our sense is a well supported group, including also the family segregates Alectoriaceae, Hypogymniaceae, Usneaceae and Anziaceae.

Lichen-forming fungi have been assumed to be more or less restricted to the surface of the substrate on which they grow, Conclusive identification of hyphae or an assessment of the fungal diversity inside lichen-covered rock has not been possible using methods based on direct observation. We circumvented this problem by using a DNA sequencing approach. Cores were drilled from a Devonian arcosic sandstone rock harboring the crustose lichen Ophioparma ventosa (L.) Norman on the surface. The cores were cut vertically, and DNA was extracted from the pulverized rock slices. A series of polymerase chain reactions using fungal-specific primers as well as Ophioparma ventosa specific primers were employed to amplify the internal transcribed spacer region of the nuclear ribosomal DNA. The results show that hyphae of O. ventosa penetrate approximately 10-12 mm into the rock. Consequently, the hyphal layer formed by the lichen fungus inside the rock could be 7-12 times as thick as the symbiotic thallus at the surface of the rock. In addition, eight non-lichenized fungal taxa and five that could not be identified to species were encountered. One fungal species in the order Helotiales occurs in six of the eight cores. The significance of these results to the colonization and weathering of rock by lichenized fungi is discussed.

Bacidia caesiovirens S. Ekman & Holien sp. nov. is described from Norway, and the British Isles. It occurs on trunks of deciduous trees in coastal forests. This species is usually sterile and is characterized by its conspicuously blue-green, granular thallus, which contains a bluish, N+ purple pigment. It is closely related to B. biatorina (Körb.) Vain., but differs from this species in a number of features apart from thallus pigmentation: the thallus granules are smaller, the amount of hyphal projections from the surface of the thallus granules is larger, and the internal pigmentation of the apothecia is different. Bacidia auerswaldii (Hepp ex Stizenb.) Mig. has a similarly granular thallus but differs from B. biatorina and B. caesiovirens in having wide, fusiform spores and smaller thallus granules. Bacidia absistens (Nyl.) Arnold has an internal apothecium pigmentation similar to B. caesiovirens, but differs in having a smooth thallus, and an abundance of minute crystals in the excipulum.

Many lichen fungi form symbioses with filamentous Nostoc cyanobacteria, which cause the lichen to swell and become extremely gelatinous when moist. Within the Lecanoromycetes, such gelatinous lichens are today mainly classified in the Collemataceae (Peltigerales, Ascomycota). We performed Bayesian MCMC, maximum likelihood, and maximum parsimony analyses of three independent markers (mtSSU rDNA, nuLSU rDNA, and RPB1), to improve our understanding of the phylogeny and classification in the Peltigerales, as well as the evolution of morphological characters that have been used for classification purposes in this group. The Collemataceae and the non-gelatinous Pannariaceae are paraphyletic but can be re-circumscribed as monophyletic if Leciophysma, Physma, Ramalodium and Staurolemma are transferred to the Pannariaceae. The gelatinous taxa transferred to the Pannariaceae deviate from other Collemataceae in having simple ascospores, and several also have a ring-shaped exciple as in other Pannariaceae, rather than the disc-shaped exciple found in the typical Collemataceae. Both Collema and Leptogium are non-monophyletic. The re-circumscribed Collemataceae shares a distinct ascus type with the sister group Placynthiaceae and the Coccocarpiaceae, whereas Pannariaceae includes a variety of structures. All Pannariaceae have one-celled ascospores, whereas all Collemataceae have two- or multi-celled spores. Reconstructions of the number of character state transformations in exciple structure, thallus gelatinosity, and ascus apex structure indicate that the number of transformations is distinctly higher than the minimum possible. Most state transformations in the exciple took place from a ring-shaped to a disc-shaped exciple. Depending on the reconstruction method, most or all transformations in thallus structure took place from a non-gelatinous to a gelatinous thallus. Gains and losses of internal structures in the ascus apex account for all or a vast majority of the number of transformations in the ascus, whereas direct transformations between asci with internal structures appear to have been rare.

This paper aims to clarify taxonomy and nomenclature of the lichen Bacidia coprodes and its closest relatives in Europe and North America. Bacidia coprodes as treated in recent works was found to consist of three distinct species in Europe and North America, the correct names of which are B. coprodes ( s. str. ), B. notarisiana , and B. granosa . In addition, a fourth species, B. inornata (Nyl.) Blomb. & Forssell, was found to be closely related to this group of lichens and is also treated here. The four species can be separated by apothecial and pycnidial pigmentation as well as conidium shape and size. They are closely related to species currently treated in Bacidia , Toninia , or Arthrosporum . Six basionyms, viz. Bilimbia coprodes , Patellaria salevensis , Bilimbia subtrachona , Lecidea chytrina , L. chytrina var. hormospora , and L. inornata , are lectotypified. Detailed descriptions and a determination key are provided.

Megalaria brodoana S. Ekman & Tensberg sp. nov. (Lecanorales, lichenized Asco- mycotina) is described from western North America (British Columbia, Washington, and Oregon). It is recognized by the following characteristics: a more or less granular, often bluish thallus; dark purplish brown to black, thick-margined apothecia; a greenish (K+ pure green, N+ purple-red) pigment in the epithecium; a brown-red to brown-purple (K+ red to purple-red, N+ orange-red) pigment in the proper exciple, hypothecium, and lower part of hymenium; a pure brown (K-, N-) pigment in the hypothecium; and one-septate, thick-walled, widely ellipsoid to subglobose spores. Megalaria brodoana occurs in humid, coastal forests along with a high number of oceanic lichens. The main phorophyte is Alnus rubra. The circumscription of the genus Megalaria is discussed. The combinations Megalaria columbiana (G. Merr.) S. Ekman and M. albocincta (Degel.) Tensberg are made.