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Lichens are the archetypal symbiosis and the one for which the term was coined. Although application of shotgun sequencing techniques has shown that many lichen symbioses can harbour more symbionts than the canonically recognized fungus and photobiont, no global census of lichen organismal composition has been undertaken. Here, we analyze the genome content of 437 lichen metagenomes from six continents, and show that four bacterial lineages occur in the majority of lichen symbioses, at a frequency on par with algal photobionts. A single bacterial genus,Lichenihabitans, occurs in nearly one-third of all lichens sampled. Genome annotations from the most common lichen bacterial symbionts suggest they are aerobic anoxygenic photoheterotrophs and produce essential vitamins, but do not fix nitrogen. We also detected secondary basidiomycete symbionts in about two-thirds of analyzed metagenomes. Our survey suggests a core set of four to seven microbial symbionts are involved in forming and maintaining lichen symbioses.

Lichen fungi live in a symbiotic association with unicellular phototrophs and most have no known aposymbiotic stage. A recent study in Molecular Ecology postulated that some of them have lost mitochondrial oxidative phosphorylation and rely on their algal partners for ATP. This claim originated from an apparent lack of ATP9, a gene encoding one subunit of ATP synthase, from a few mitochondrial genomes. Here, we show that while these fungi indeed have lost the mitochondrial ATP9, each retain a nuclear copy of this gene. Our analysis reaffirms that lichen fungi produce their own ATP.

An examination of collections from Japan has increased the number of Brianaria and Micarea species known from that country from eight to 19, including one new species, M. rubioides Coppins (also from Malaysia and the Philippines). Eleven species are reported as new to Japan (M. botryoides (Nyl.) Coppins, M. denigrata (Fr.) Hedl., M. erratica (Körb.) Hertel et al., M. hedlundii Coppins, M. lithinella (Nyl.) Hedl., M. micrococca (Körb.) Gams ex Coppins and M. misella (Nyl.) Hedl.) or new to Asia: M. byssacea (Th. Fr.) Czarnota et al., M. deminuta Coppins and M. xanthonica Coppins & Tønsberg (new to Asia; Japan); M. nitschkeana (J. Lahm ex Rabenh.) Harm. (new to Asia; South Korea). The presence of Micarea prasina s. str. from Japan needs to be confirmed; no collection was found in this study. Additional collections from South Korea and Sri Lanka are also reported, including the new species M. ceylanica Coppins from Sri Lanka. The identity of M. synotheoides (Nyl.) Coppins, originally described from Japan, has been resolved, resulting in the renaming of Western European material, previously under that name, as M. longispora Coppins. Micarea coreana Lőkös et al. is reported here as a synonym of M. erratica. The type of Lecidea inopinula Nyl. requires the new combination Micarea inopinula (Nyl.) Coppins & T. Sprib. to replace Micarea prasinella (Jatta) I. M. Lamb.

Mountain caribou, a behaviourally and genetically distinct set of ecotypes of the Woodland caribou (Rangifer tarandus caribou) restricted to the mountains of western Canada, have undergone severe population declines in recent decades. Although a broad consensus exists that the ultimate driver of these declines has been the reduction of habitat upon which mountain caribou depend, research and policy attention has increasingly focused on predation. Recently, Serrouya et al. (Proc Nat Acad Sci USA 116:6181–6186, 2019) analysed population dynamics data from 18 subpopulations in British Columbia and Alberta, Canada, subject to different treatments and ‘controls’, and concluded that lethal wolf control and maternal caribou penning provide the most effective ways to stabilize population declines. Here we show that this inference was based on an unbalanced analytical approach that omitted a null scenario, excluded potentially confounding variables and employed irreproducible habitat alteration metrics. Our reanalysis of available data shows that ecotype identity is a better predictor of population trends than any adaptive management treatments considered by Serrouya et al. Disparate behavioural characteristics and responses to industrial disturbance among ecotypes suggest it may be incorrect to assume that adaptive management strategies that might benefit one ecotype are transferable to another.

Ascomycota account for about two-thirds of named fungal species.1 Over 98% of known Ascomycota belong to the Pezizomycotina, including many economically important species as well as diverse pathogens, decomposers, and mutualistic symbionts.2 Our understanding of Pezizomycotina evolution has until now been based on sampling traditionally well-defined taxonomic classes.3,4,5 However, considerable diversity exists in undersampled and uncultured, putatively early-diverging lineages, and the effect of these on evolutionary models has seldom been tested. We obtained genomes from 30 putative early-diverging lineages not included in recent phylogenomic analyses and analyzed these together with 451 genomes covering all available ascomycete genera. We show that 22 of these lineages, collectively representing over 600 species, trace back to a single origin that diverged from the common ancestor of Eurotiomycetes and Lecanoromycetes over 300 million years BP. The new clade, which we recognize as a more broadly defined Lichinomycetes, includes lichen and insect symbionts, endophytes, and putative mycorrhizae and encompasses a range of morphologies so disparate that they have recently been placed in six different taxonomic classes. To test for shared hidden features within this group, we analyzed genome content and compared gene repertoires to related groups in Ascomycota. Regardless of their lifestyle, Lichinomycetes have smaller genomes than most filamentous Ascomycota, with reduced arsenals of carbohydrate-degrading enzymes and secondary metabolite gene clusters. Our expanded genome sample resolves the relationships of numerous “orphan” ascomycetes and establishes the independent evolutionary origins of multiple mutualistic lifestyles within a single, morphologically hyperdiverse clade of fungi.

Lichens are the symbiotic outcomes of open, interspecies relationships, central to which are a fungus and a phototroph, typically an alga and/or cyanobacterium. The evolutionary processes that led to the global success of lichens are poorly understood. In this review, we explore the goods and services exchange between fungus and phototroph and how this propelled the success of both symbiont and symbiosis. Lichen fungal symbionts count among the only filamentous fungi that expose most of their mycelium to an aerial environment. Phototrophs export carbohydrates to the fungus, which converts them to specific polyols. Experimental evidence suggests that polyols are not only growth and respiratory substrates but also play a role in anhydrobiosis, the capacity to survive desiccation. We propose that this dual functionality is pivotal to the evolution of fungal symbionts, enabling persistence in environments otherwise hostile to fungi while simultaneously imposing costs on growth. Phototrophs, in turn, benefit from fungal protection from herbivory and light stress, while appearing to exert leverage over fungal sex and morphogenesis. Combined with the recently recognized habit of symbionts to occur in multiple symbioses, this creates the conditions for a multiplayer marketplace of rewards and penalties that could drive symbiont selection and lichen diversification.

Since the advent of molecular taxonomy, numerous lichen-forming fungi with homoiomerous thalli initially classified in the family Collemataceae Zenker have been transferred to other families, highlighting the extent of morphological convergence within Lecanoromycetes O. E. Erikss. & Winka. While the higher level classification of these fungi might be clarified by such transfers, numerous specific and generic classifications remain to be addressed. We examined the relationships within the broadly circumscribed genus Arctomia Th. Fr., which has been the recipient of several transfers from Collemataceae. We demonstrated that Arctomia insignis (P. M. Jørg. & Tønsberg) Ertz does not belong to Arctomia s. str. but forms a strong monophyletic group with Gabura fascicularis (L.) P. M. Jørg. We also confirmed that Arctomia borbonica Magain & Sérus. and the closely related Arctomia insignis represent two species. We formally transferred A. insignis and A. borbonica to the genus Gabura Adans. and introduced two new combinations: Gabura insignis and Gabura borbonica. We reported Gabura insignis from Europe (Scotland and Ireland) for the first time. While material from Europe and North America is genetically almost identical, specimens from Madagascar, South Africa and Reunion Island belong to three distinct phylogenetic lineages, all of which are present in the latter area and may represent distinct species. In its current circumscription, the genus Gabura may contain up to six species, whereas Arctomia s. str. includes only two species (A. delicatula Th. Fr. and A. teretiuscula P. M. Jørg.). The Gabura insignis group is shown to have an unexpectedly large, subcosmopolitan distribution. With the extended sampling from Arctomiaceae Th. Fr., the placement of Steinera sorediata P. James & Henssen in the genus Steinera Zahlbr. is confirmed and the presence of a new Steinera species from Chile is highlighted.

Lichen symbioses are generally thought to be stabilized by the transfer of fixed carbon compounds from a photosynthesizing unicellular symbiont to a fungus. In other fungal symbioses, carbohydrate subsidies correlate with genomic reductions in the number of genes for plant cell wall-degrading enzymes (PCWDEs), but whether this is the case with lichen fungal symbionts (LFSs) is unknown. We predicted genes encoding carbohydrate-active enzymes (CAZymes) and sugar transporters in 17 existing and 29 newly sequenced genomes from across the class Lecanoromycetes, the largest extant clade of LFSs. Despite possessing lower mean numbers of PCWDE genes compared to non-symbiont Ascomycota, all LFS genomes possessed a robust suite of predicted PCWDEs. The largest CAZyme gene numbers, on par with model species such as Penicillium, were retained in genomes from the subclass Ostropomycetidae, which are found in crust lichens with highly specific ecologies. The lowest numbers were in the subclass Lecanoromycetidae, which are symbionts of many generalist macrolichens. Our results suggest that association with phototroph symbionts does not in itself result in functional loss of PCWDEs and that PCWDE losses may have been driven by adaptive processes within the evolution of specific LFS lineages. The inferred capability of some LFSs to access a wide range of carbohydrates suggests that some lichen symbioses may augment fixed CO2 with carbon from external sources.SignificanceLichen symbioses are considered self-contained autotrophic systems in which the total carbon economy is the sum of phototroph-fixed CO2, supplied to a fungus as sugars. In other fungal-plant symbioses, such as mycorrhizae, plant-derived sugar subsidies are associated with loss of plant cell wall-degrading enzymes (PCWDEs). We compared PCWDE inventories in 46 genomes from the largest group of lichen fungal symbionts (LFSs) with non-symbionts from across Ascomycota. We found that despite lower overall gene numbers, all LFSs retain PCWDEs, and some possess gene numbers and functional diversity on par with non-symbionts. Our results suggest that association with a phototroph does not necessarily result in PCWDE loss, and some lichens may obtain carbon from sources other than CO2 fixation.

Konoreva, L. A., Chesnokov, S. V., Stepanchikova, I. S., Spribille, T., Bjork, C. & Williston, P. 2021. Nine Micarea species new to Canada including five species new to North America. – Herzogia 34: 18 –37.Nine species of the genus Micarea are reported for the first time from Canada including five (M. contexta, M. fallax, M. pseudomicrococca, M. pusilla, M. tomentosa) which are new to North America. Most of them are part of the Micarea prasina complex. Canadian records of further rare or taxonomically challenging species (M. botryoides, M. inopinula, M. prasina s. str.) are revised. The Canadian records are discussed in the context of their wider distribution, and differences to related or morphologically similar species are indicated.

Basidiomycete yeasts have recently been reported as stably associated secondary fungal symbionts of many lichens, but their role in the symbiosis remains unknown. Attempts to sequence their genomes have been hampered both by the inability to culture them and their low abundance in the lichen thallus alongside two dominant eukaryotes (an ascomycete fungus and chlorophyte alga). Using the lichen Alectoria sarmentosa, we selectively dissolved the cortex layer in which secondary fungal symbionts are embedded to enrich yeast cell abundance and sequenced DNA from the resulting slurries as well as bulk lichen thallus. In addition to yielding a near-complete genome of the filamentous ascomycete using both methods, metagenomes from cortex slurries yielded a 36- to 84-fold increase in coverage and near-complete genomes for two basidiomycete species, members of the classes Cystobasidiomycetes and Tremellomycetes. The ascomycete possesses the largest gene repertoire of the three. It is enriched in proteases often associated with pathogenicity and harbors the majority of predicted secondary metabolite clusters. The basidiomycete genomes possess ∼35% fewer predicted genes than the ascomycete and have reduced secretomes even compared with close relatives, while exhibiting signs of nutrient limitation and scavenging. Furthermore, both basidiomycetes are enriched in genes coding for enzymes producing secreted acidic polysaccharides, representing a potential contribution to the shared extracellular matrix. All three fungi retain genes involved in dimorphic switching, despite the ascomycete not being known to possess a yeast stage. The basidiomycete genomes are an important new resource for exploration of lifestyle and function in fungal-fungal interactions in lichen symbioses.

Lichen fungi live in a symbiotic association with unicellular phototrophs and have no known aposymbiotic stage. A recent study postulated that some of them have lost mitochondrial oxidative phosphorylation and rely on their algal partners for ATP. This claim originated from an apparent lack of ATP9, a gene encoding one subunit of ATP synthase, from a few mitochondrial genomes. Here we show that while these fungi indeed have lost the mitochondrial ATP9, each retain a nuclear copy of this gene. Our analysis reaffirms that lichen fungi produce their own ATP.

Basidiomycete yeasts have recently been reported as stably associated secondary fungal symbionts of many lichens, but their role in the symbiosis remains unknown. Attempts to sequence their genomes have been hampered both by the inability to culture them and their low abundance in the lichen thallus alongside two dominant eukaryotes (an ascomycete fungus and chlorophyte alga). Using the lichen Alectoria sarmentosa, we selectively dissolved the cortex layer in which secondary fungal symbionts are embedded to enrich yeast cell abundance and sequenced DNA from the resulting slurries as well as bulk lichen thallus. In addition to yielding a near-complete genome of the filamentous ascomycete using both methods, metagenomes from cortex slurries yielded a 36- to 84-fold increase in coverage and near-complete genomes for two basidiomycete species, members of the classes Cystobasidiomycetes and Tremellomycetes. The ascomycete possesses the largest gene repertoire of the three. It is enriched in proteases often associated with pathogenicity and harbors the majority of predicted secondary metabolite clusters. The basidiomycete genomes possess ∼35% fewer predicted genes than the ascomycete and have reduced secretomes even compared with close relatives, while exhibiting signs of nutrient limitation and scavenging. Furthermore, both basidiomycetes are enriched in genes coding for enzymes producing secreted acidic polysaccharides, representing a potential contribution to the shared extracellular matrix. All three fungi retain genes involved in dimorphic switching, despite the ascomycete not being known to possess a yeast stage. The basidiomycete genomes are an important new resource for exploration of lifestyle and function in fungal–fungal interactions in lichen symbioses.

The study of the reproductive biology of lichen fungal symbionts has been traditionally challenging due to their complex and symbiotic lifestyles. Against the common belief of haploidy, a recent genomic study found a triploid-like signal inLetharia. Here, we used genomic data from a pure culture and from thalli, together with a PCR survey of the MAT locus, to infer the genome organization and reproduction inLetharia. We found that the read count variation in the fourLethariaspecimens, including the pure culture derived from a single sexual spore ofL. lupina, is consistent with haploidy. By contrast, theL. lupinaread counts from a thallus’ metagenome are triploid-like. Characterization of the mating-type locus revealed a conserved heterothallic configuration across the genus, along with auxiliary genes that we identified. We found that the mating-type distributions are balanced in North America forL. vulpinaandL. lupina, suggesting widespread sexual reproduction, but highly skewed in Europe forL. vulpina, consistent with predominant asexuality. Taken together, we propose thatLethariafungi are heterothallic and typically haploid, and provide evidence that triploid-like individuals are rare hybrids betweenL. lupinaand an unknownLetharialineage, reconciling classic systematic and genetic studies with recent genomic observations.

The lichen, to which the name Lecidea lichenicola is found to have been misapplied, was first described from England and is an extreme specialist of chalk pebbles. It has long been known that it is not closely related to Lecidea in the strict sense, but its true evolutionary relationships have been unknown. Here we use metagenome-assembled genome data to place this fungus in a six-locus phylogeny of Ascomycota, and find strong support for its placement in the class Lichinomycetes. Multiple gene trees using existing data from Lichinomycetes support its further placement within the family Lichinaceae. Based on a revision of types and original descriptions, we conclude that the earliest name for this species is Lecidea obsoleta (syn. Thrombium cretaceum). We neotypify that name by a modern collection and accommodate it in the new genus Watsoniomyces. Type and other original material of L. lichenicola (syn. Discocera lichenicola) was re-examined and found not to be on chalk and to represent a different lichen, Trapelia glebulosa. Watsoniomyces is the first described member of Lichinomycetes with an endolithic thallus.

Fungi critically impact the health and function of global ecosystems and economies. In Canada, fungal researchers often work within silos defined by subdiscipline and institutional type, complicating the collaborations necessary to understand the impacts fungi have on the environment, economy, and plant and animal health. Here, we announce the establishment of the Canadian Fungal Research Network (CanFunNet, https://fungalresearch.ca ), whose mission is to strengthen and promote fungal research in Canada by facilitating dialogue among scientists. We summarize the challenges and opportunities for Canadian fungal research that were discussed at CanFunNet’s inaugural meeting in 2019, and identify 4 priorities for our community: (i) increasing collaboration among scientists, (ii) studying diversity in the context of ecological disturbance, (iii) preserving culture collections in the absence of sustained funding, and (iv) leveraging diverse expertise to attract trainees. We have gathered additional information to support our recommendations, including a survey identifying underrepresentation of fungal-related courses at Canadian universities, a list of Canadian fungaria and culture collections, and a case study of a human fungal pathogen outbreak. We anticipate that these discussions will help prioritize fungal research in Canada, and we welcome all researchers to join this nationwide effort to enhance knowledge dissemination and funding advocacy.

Nomenclatural type definitions are one of the most important concepts in biological nomenclature. Being physical objects that can be re-studied by other researchers, types permanently link taxonomy (an artificial agreement to classify biological diversity) with nomenclature (an artificial agreement to name biological diversity). Two proposals to amend the International Code of Nomenclature for algae, fungi, and plants (ICN), allowing DNA sequences alone (of any region and extent) to serve as types of taxon names for voucherless fungi (mainly putative taxa from environmental DNA sequences), have been submitted to be voted on at the 11th International Mycological Congress (Puerto Rico, July 2018). We consider various genetic processes affecting the distribution of alleles among taxa and find that alleles may not consistently and uniquely represent the species within which they are contained. Should the proposals be accepted, the meaning of nomenclatural types would change in a fundamental way from physical objects as sources of data to the data themselves. Such changes are conducive to irreproducible science, the potential typification on artefactual data, and massive creation of names with low information content, ultimately causing nomenclatural instability and unnecessary work for future researchers that would stall future explorations of fungal diversity. We conclude that the acceptance of DNA sequences alone as types of names of taxa, under the terms used in the current proposals, is unnecessary and would not solve the problem of naming putative taxa known only from DNA sequences in a scientifically defensible way. As an alternative, we highlight the use of formulas for naming putative taxa (candidate taxa) that do not require any modification of the ICN. Publisher’s Note A first version of this text was prepared by the first eight authors and the last one, given here. The other listed co-authors in the article PDF support the content, and their actual contributions varied from only support to additions that substantially improved the content. The full details of all co-authors, with their affiliations, are included in Supplementary Table 1 after p.175 of the article for reasons of clarity and space. Slavomír Adamčík Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Teuvo Ahti Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland M. Catherine Aime Purdue University, 915 W. State St., West Lafayette, Indiana 47907, U.S.A. A. Martyn Ainsworth Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom László Albert Hungarian Mycological Society, 1087 Könyves Kálmán krt. 40, Budapest, Hungary Edgardo Albertó Instituto de Investigaciones Biotecnológicas-Instituto Tecnológico de Chascomús, Universidad Nacional de San Martin-Consejo Nacional de Investigaciones Científicas y Técnicas, Buenos Aires, Argentina Alberto Altés García Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Dmitry Ageev SIGNATEC Ltd., 630090, Novosibirsk, Akademgorodok (Novosibirsk Scientific Center), Inzhenernaya str., 22, Russia Reinhard Agerer Ludwig-Maximilians-Universität München, Menzinger Str. 67, 80638 München, Germany Begona Aguirre-Hudson Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom Joe Ammirati University of Washington, Seattle, Washington 98195-1800, U.S.A. Harry Andersson Eichhahnweg 29a, 38108 Braunschweig, Germany Claudio Angelini Jardín Botánico Nacional Dr. Rafael Ma. Moscoso, Apartado 21-9, Santo Domingo, Dominican Republic Vladimír Antonín Moravian Museum, Zeny trh 6, 659 37 Brno, Czech Republic Takayuki Aoki Genetic Resources Center, National Agriculture and Food Research Organization, 2-1-2 Kannondai, Tsukuba, Ibaraki 305-8602, Japan André Aptroot ABL Herbarium, G.v.d.Veenstraat 107, 3762 XK Soest, The Netherlands Didier Argaud 40 rue du Justemont, 57290 Fameck, France Blanca Imelda Arguello Sosa Instituto Tecnológico de Ciudad Victoria, Tecnológico Nacional de México, Ciudad Victoria, Tamaulipas, Mexico Arne Aronsen Torødveien 54, 3135 Torød, Norway Ulf Arup Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Bita Asgari Iranian Research Institute of Plant Protection, Agricultural Research, Education and Extension Organization, Tehran, Iran Boris Assyov Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Str., 1113 Sofia, Bulgaria Violeta Atienza Facultad de Ciencias Biológicas, Universitat de València, C/Dr Moliner 50, 46100, Burjasot, Valencia, Spain Ditte Bandini Panoramastr 47, 69257 Wiesenbach, Germany João Luís Baptista-Ferreira Instituto de Biossistemas e Ciências Integrativas, Faculdade de Ciências da Universidade de Lisboa, 1749-016 Lisboa, Portugal Hans-Otto Baral Blaihofstr. 42, 72074 Tübingen, Germany Tim Baroni The State University of New York, 340 Bowers Hall, P.O. Box 2000, Cortland, New York 13045, U.S.A. Robert Weingart Barreto Universidade Federal de Viçosa, 36570-000, Viçosa, Minas Gerais, Brazil Henry Beker (1) Royal Holloway, University of London, United Kingdom; (2) Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Ann Bell 45 Gurney Road, Lower Hutt, New Zealand Jean-Michel Bellanger CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry Montpellier, EPHE, INSERM, 1919 Route de Mende, 34293 Montpellier Cédex 5, France Francesco Bellù Naturmusem of Bolzano, CP 104, 39100, Bolzano, Italy Martin Bemmann Kleingemünderstraße 111, 69118 Heidelberg, Germany Mika Bendiksby NTNU, University Museum, Norwegian University of Science and Technology, 7491 Trondheim, Norway Egil Bendiksen Norwegian Institute for Nature Research, Gaustadalleen 21, 0349 Oslo, Norway Katriina Bendiksen Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Lajos Benedek Szent Istvan University, Hungary Anna Bérešová-Guttová Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Franz Berger University of Salzburg, Salzburg, Austria Reinhard Berndt Herbaria Z+ZT, ETH Zürich, CHN D37, Universitätstr. 16, 8092 Zürich, Switzerland Annarosa Bernicchia Via A. Guidotti 39, 40134 Bologna, Italy Alona Yu. Biketova Institute of Biochemistry, BRC-HAS, 6726 Szeged, Temesvari krt. 62, 6726 Szeged, Hungary Enrico Bizio Società Veneziana di Micologia, Società Veneziana di Scienze Naturali, Fontego dei Turchi, Santa Croce 1730, 30135 Venice, Italy Curtis Bjork UBC Herbarium, Beaty Biodiversity Museum, University of British Columbia, Canada Teun Boekhout (1) Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands; (2) Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Amsterdam, The Netherlands David Boertmann Aarhus University, Frederiksborgvej 399, 4000 Roskilde, Denmark Tanja Böhning AG Geobotanik Schleswig-Holstein & Hamburg, c/o University of Kiel, Olshausenstraße 75, 24098 Kiel, Germany Florent Boittin Ascomycete.org, 36 rue de la Garde, 69005 Lyon, France Carlos G. Boluda Conservatoire et Jardin botaniques de la Ville de Genève, 1292 Genève, Switzerland Menno W. Boomsluiter T.v.Lohuizenstraat 34, 8172xl, Vaassen, The Netherlands Jan Borovička Institute of Geology, Czech Academy of Sciences, Rozvojova 269, 165 00 Prague 6, Czech Republic Tor Erik Brandrud Norwegian Institute for Nature Research, Gaustadalleen 21, 0349 Oslo, Norway Uwe Braun Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik, und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle, Germany Irwin Brodo Canadian Museum of Nature, 240 McLeod Street, Ottawa, Ontario, Canada Tatiana Bulyonkova A.P. Ershov Institute of Informatics Systems, Russian Academy of Sciences, Siberian Branch, 6 Acad. Lavrentjev pr., Novosibirsk 630090, Russia Harold H. Burdsall Jr. Fungal & Decay Diagnostics, LLC, 9350 Union Valley Road, Black Earth, Wisconsin 53515, U.S.A. Bart Buyck Muséum National d’Histoire Naturelle, CP 39, ISYEB, UMR 7205 CNRS MNHN UPMC EPHE, 12 rue Buffon, 75005 Paris, France Ana Rosa Burgaz Facultad de Ciencias Biológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain Vicent Calatayud Fundación CEAM, c/ Charles R. Darwin, 14, Parque Tecnológico, 46980 Paterna, Valencia, Spain Philippe Callac INRA, MycSA, CS 20032, 33882 Villenave d’Ornon, France Emanuele Campo Associazione Micologica Bresadola, Via Alessandro Volta 46, 38123 Trento, Italy Massimo Candusso Via Ottone Primo 90, 17021, Alassio, Savona, Italy Brigitte Capoen Queffioec, rue de Saint Gonval, 22710 Penvenan, France Joaquim Carbó Roser, 60, 17257 Torroella de Montgrí, Girona, Spain Matteo Carbone Via Don Luigi Sturzo 173 16148 Genova, Italy Rafael F. Castañeda-Ruiz Instituto de Investigaciones Fundamentales en Agricultura, Tropical ‘Alejandro de Humboldt’, OSDE, Grupo Agrícola, Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana 17200, Cuba Michael A. Castellano USDA, Forest Service, Northern Research Station, Corvallis, Oregon 97330, U.S.A. Jie Chen Mae Fah Luang University, Chang Wat Chiang Rai 57100, Thailand Philippe Clerc Conservatoire et Jardin botaniques de la Ville de Genève, 1292 Genève, Switzerland Giovanni Consiglio Via C. Ronzani 61, 40033 Casalecchio Bologna, Italy Gilles Corriol National Botanical Conservatory for Pyrenees and Midi-Pyrénées Region of France and BBF Herbarium, Vallon de Salut. B.P. 315. 65203 Bagnères-de-Bigorre, France Régis Courtecuisse Université Lille, Fac. Pharma. Lille, EA4483 IMPECS, 59000 Lille, France Ana Crespo Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain Cathy Cripps Plant Sciences & Plant Pathology, 119 Plant Biosciences Building, Montana State University, Bozeman, Montana 59717, U.S.A. Pedro W. Crous Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Gladstone Alves da Silva Universidade Federal de Pernambuco, Centro de Biociências, Avenida da Engenharia, S/N, Cidade Universitária, Recife, Pernambuco, Brazil Meiriele da Silva Universidade Federal de Viçosa, 36570-000, Viçosa, Minas Gerais, Brazil Marjo Dam Hooischelf 13, 6581 SL Malden, The Netherlands Nico Dam Hooischelf 13, 6581 SL Malden, The Netherlands Frank Dämmrich The Bavarian Natural History Collections (SNSB Munich), Menzinger Strasse 71, 80638, München, Germany Kanad Das Botanical Survey of India, Cryptogamic Unit, P.O. Botanic Garden, Howrah 711103, W.B., India Linda Davies Centre for Environmental Policy, Imperial College London, SW7 2AZ, United Kingdom Eske De Crop Ghent University K.L. Ledeganckstraat 35, 9000 Ghent, Belgium Andre De Kesel Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Ruben De Lange Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium Bárbara De Madrignac Bonzi Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Thomas Edison E. dela Cruz University of Santo Tomas, Espana 1008 Manila, Philippines Lynn Delgat Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium Vincent Demoulin Institut de Botanique, B.22, Université de Liège, 4000 Liège I, Belgium Dennis E. Desjardin HD Thiers Herbarium (SFSU), San Francisco State University, 1600 Holloway Ave, San Francisco, California 94132, U.S.A. Paul Diederich Musée national d’histoire naturelle, 25 rue Münster, 2160 Luxembourg, Luxembourg Bálint Dima (1) Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; (2) Viikki Plant Science Centre, University of Helsinki, P.O. Box 65, 00014 Helsinki, Finland Maria Martha Dios Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Catamarca, Av Belgrano 300, 4700 San Fernando del Valle de Catamarca, Argentina Pradeep Kumar Divakar Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain Clovis Douanla-Meli Julius Kühn-Institut, Federal Research Centre for Cultivated Plants, Institute for National and International Plant Health, Messeweg 11-12, 38104 Braunschweig, Germany Brian Douglas Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom Elisandro Ricardo Drechsler-Santos Universidade Federal de Santa Catarina, Campus Universitário Reitor João David Ferreira Lima, Trindade, Florianópolis, Santa Catarina CEP 88040-900, Brazil Paul S. Dyer School of Life Sciences, University of Nottingham, University Park, Nottingham NG7 2RD, United Kingdom Ursula Eberhardt Abt. Botanik, Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany Damien Ertz Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Fernando Esteve-Raventós Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Javier Angel Etayo Salazar Navarro Villoslada 16, 3º dcha., 31003 Pamplona, Navarra, Spain Vera Evenson Sam Mitchel Herbarium of Fungi, Denver Botanic Gardens, 1007 York Street, Denver, Colorado 80206, U.S.A. Guillaume Eyssartier Muséum national d’histoire naturelle, Jardin des plantes, 57 rue Cuvier, 75005 Paris, France Edit Farkas Institute of Ecology and Botany, MTA Centre for Ecological Research, 2163 Vácrátót, Hungary Alain Favre Fédération Mycologique et Botanique Dauphiné Savoie, Le Prieuré, 144 Place de l’Eglise, 74320 Sevrier, France Anna G. Fedosova Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Mario Filippa Regione Monsarinero 36, 14041 Agliano Terme, Italy Péter Finy 8000 Székesfehérvár, Zsombolyai u. 56, Hungary Adam Flakus W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Simón Fos Facultad de Ciencias Biológicas, Universitat de València, C/Dr Moliner 50, 46100, Burjasot, Valencia, Spain Jacques Fournier Las Muros, F. 09420 Rimont, France André Fraiture Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Paolo Franchi Associazione Micologica Bresadola, Via Alessandro Volta 46, 38123 Trento, Italy Ana Esperanza Franco Molano Escuela de Microbiología, Universidad de Antioquia, AA1226, Fundación Biodiversa Colombia, Medellín, Colombia Gernot Friebes Centre of Natural History, Botany & Mycology, Universalmuseum Joanneum, Weinzöttlstraße 16, 8045 Graz, Austria Andreas Frisch NTNU, University Museum, Norwegian University of Science and Technology, 7491 Trondheim, Norway Alan Fryday Michigan State University, East Lansing, Michigan 48824, U.S.A. Giuliana Furci The Fungi Foundation, Paseo Bulnes 79 of. 112A, Santiago, Chile Ricardo Galán Márquez Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Matteo Garbelotto University of California, 130 Mulford Hall #3114 Berkeley, California 94720, U.S.A. Joaquina Maria Garcia-Martin Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Mónica A. García Otálora Herbaria Z+ZT, ETH Zürich, CHN D37, Universitätstr. 16, 8092 Zürich, Switzerland Dania García Sánchez Universitat Rovira i Virgili, C/ Sant Llorenç 21, 43201 Reus, Tarragona, Spain Alain Gardiennet 14 rue Roulette, 21260 Véronnes, France Sigisfredo Garnica Instituto de Bioquímica y Microbiología, Universidad Austral de Chile, Isla Teja Campus, Casilla 567, Valdivia, Chile Isaac Garrido Benavent Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Genevieve Gates Tasmanian Institute of Agriculture, Private Bag 54, Hobart, Tasmania 7001, Australia Alice da Cruz Lima Gerlach Conservatoire et Jardin Botaniques de la ville de Genève, Genève, Switzerland Masoomeh Ghobad-Nejhad Iranian Research Organization for Science and Technology, P.O. Box 15815-3538, Tehran 15819, Iran Tatiana B. Gibertoni Universidade Federal de Pernambuco, Centro de Biociências, Avenida da Engenharia, S/N, Cidade Universitária, Recife, Pernambuco, Brazil Tine Grebenc Slovenian Forestry Institute, Vecna pot 2, 100 Ljubljana, Slovenia Irmgard Greilhuber University of Vienna, Rennweg 14, 1030 Vienna, Austria Bella Grishkan Institute of Evolution, University of Haifa, Aba Khoushi Ave. 199, Mt. Carmel, Haifa 3498838, Israel Johannes Z. Groenewald Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Martin Grube Institute of Biology, University of Graz, Holteiasse 6, 8010 Graz, Austria Gérald Gruhn Office National des Forêts, 2 Avenue de Saint-Mandé, 75570 Paris Cedex 12, France Cécile Gueidan CSIRO — Australian National Herbarium, Clunies Ross Street, Canberra ACT 2601, Australia Gro Gulden Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Luis FP Gusmão Universidade Estadual de Feira de Santana, Av. Transnordestina, s/n, Bairro Novo Horizonte, CEP:44036-900, Feira de Santana, Bahia, Brazil Josef Hafellner Institute of Biology, University of Graz, Holteiasse 6, 8010 Graz, Austria Michel Hairaud 2 Impasse des Marronniers, 79360 Poivendre de Marigny, France Marek Halama Museum of Natural History, Wrocław University, ul. H. Sienkiewicza 5, 50-335 Wrocław, Poland Nils Hallenberg University of Gothenburg, Box 461, 40530 Göteborg, Sweden Roy E. Halling Institute of Systematic Botany, New York Botanical Garden, 2900 Southern Blvd, Bronx, New York 10458-5126, U.S.A. Karen Hansen Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Christoffer Bugge Harder Texas Tech University, Box 42122, Lubbock, Texas 79409, U.S.A. Jacob Heilmann-Clausen Natural History Museum of Denmark, Universitetsparken 15, 2100 København, Denmark Stip Helleman Sweelinck 78, 5831KT Boxmeer, The Netherlands Alain Henriot Mycological Society of France, 20 rue Rottembourg, 12th arrondissement, Paris, France Margarita Hernandez-Restrepo Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Raphaël Herve 24 rue des Fougères, 86550 Mignaloux-Beauvoir, France Caroline Hobart 84 Stafford Road, Sheffield, South Yorkshire S2 2SF, United Kingdom Mascha Hoffmeister Julius Kühn-Institut, Institute for Epidemiology and Pathogen Diagnostics, Braunschweig, Germany Klaus Høiland University of Oslo, P.O.Box 1066, Blindern, 0316 Oslo, Norway Jan Holec National Museum, Herbarium PRM, Cirkusová 1740, 193 00 Praha 9, Czech Republic Håkon Holien Faculty of Bioscience and Aquaculture, NORD University, P.O. Box 2501, 7729 Steinkjer, Norway Karen Hughes University of Tennessee, Knoxville, Tennessee 37996, U.S.A. Vit Hubka Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Seppo Huhtinen Herbarium TUR, Biodiversity Unit, University of Turku, 20014 Turku, Finland Boris Ivančević Natural History Museum, Njegoševa 51, P.O. Box 401, 11000 Belgrade, Serbia Marian Jagers Reelaan 13, 7522 LR Enschede, The Netherlands Walter Jaklitsch Institute of Forest Entomology, Forest Pathology and Forest Protection, University of Natural Resources and Life Sciences Vienna, Vienna, Austria AnnaElise Jansen Stationsstraat 10, 6701 AM Wageningen, the Netherlands Ruvishika S. Jayawardena Mae Fah Luang University, Chang Wat Chiang Rai 57100, Thailand Thomas Stjernegaard Jeppesen Global Biodiversity Information Facility, Universitetsparken 15, 2100 København Ø, Denmark Mikael Jeppson Lilla Håjumsgatan 4, 46135 Trollhättan, Sweden Peter Johnston Manaaki Whenua Landcare Research, Private Bag 92170, Auckland 1072, New Zealand Per Magnus Jørgensen University of Bergen, Allégaten 41, P.O. Box 7800, 5020 Bergen, Norway Ingvar Kärnefelt Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Liudmila B. Kalinina Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Gintaras Kantvilas Tasmanian Herbarium (HO), Tasmanian Museum and Art Gallery, P.O. Box 5058, UTAS LP.O., Sandy Bay, Tasmania 7005, Australia Mitko Karadelev Institute of Biology, Faculty of Natural Science and Mathematics, Ss. Cyril and Methodius University, Arhimedova 5, 1000 Skopje, Republic of Macedonia Taiga Kasuya Faculty of Risk and Crisis Management, Chiba Institute of Science, 3 Shiomi-cho, Choshi, Chiba 288-0025, Japan Ivona Kautmanová Natural History Museum, Slovak National Museum, Bratislava, Slovakia Richard W. Kerrigan RWK Research, Kittanning, Pennsylvania 16201, U.S.A. Martin Kirchmair Institut für Mikrobiologie, Universität Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria Anna Kiyashko Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Dániel G. Knapp Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary Henning Knudsen Natural History Museum of Denmark, Universitetsparken 15, 2100 København, Denmark Kerry Knudsen Faculty of Environmental Sciences, University of Life Sciences at Prague, Prague, Czech Republic Tommy Knutsson Nedra Västerstad 111, 380 62 Mörbylånga, Sweden Miroslav Kolařík Institute of Microbiology ASCR, Videnska 1083, 142 20 Prague 4, Czech Republic Urmas Kõljalg Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Alica Košuthová Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Attila Koszka Faculty of Agricultural and Environmental Sciences, Kaposvar University, 7400 Kaposvar, Hungary Heikki Kotiranta Finnish Environment Institute, P.O. Box 140, 00251 Helsinki, Finland Vera Kotkova Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Ondřej Koukol Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Jiří Kout University of West Bohemia, Faculty of Education, Klatovska 51, 306 19 Pilsen, Czech Republic Gábor M. Kovács Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary Martin Kříž Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Åsa Kruys Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Viktor Kučera Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23 Bratislava, Slovakia Linas Kudzma 37 Maple Ave. Annandale, New Jersey 08801, U.S.A. Francisco Kuhar Instituto Multidisciplinario de Biología Vegetal, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Córdoba, Casilla de Correo 495, 5000 Córdoba, Argentina Martin Kukwa Faculty of Biology, University of Gdańsk, Wita Stwosza 59, 80-308 Gdańsk, Poland T. K. Arun Kumar The Zamorin’s Guruvayurappan College, Kozhikode, Kerala 673014, India Vladimír Kunca Technical University in Zvolen, Ul. T. G. Masaryka 24, 960 53 Zvolen,Slovakia Ivana Kušan Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Thomas W. Kuyper Wageningen University & Research, Droevendaalsesteeg 4, 6708 PB Wageningen, The Netherlands Carlos Lado Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Thomas Læssøe Natural History Museum of Denmark, Universitetsparken 15, 2100 København, Denmark Patrice Lainé 123 rue Saint Antoine, 75004, Paris, France Ewald Langer University of Kassel, Heinrich-Plett-Strasse 40, 34132 Kassel, Germany Ellen Larsson University of Gothenburg, Box 461, 40530 Göteborg, Sweden Karl-Henrik Larsson Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Gary Laursen Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, Alaska 99775-7000, U.S.A. Christian Lechat Ascofrance, 64 route de Chizé, 79360 Villiers en Bois, France Serena Lee Herbarium Singapore Botanic Gardens, National Parks Board, Singapore James C. Lendemer (1) Institute of Systematic Botany, New York Botanical Garden, 2900 Southern Blvd, Bronx, New York 10458-5126, U.S.A.; (2) Graduate Center, City University of New York, 365 5th Ave, New York, New York 10016, U.S.A. Laura Levin University of Buenos Aires, Junin 956, 1113, Buenos Aires, Argentina Uwe Lindemann Landesmuseum für Naturkunde, Münster, Germany Håkan Lindström Östansjö 150, 840 64 Kälarne, Sweden Xingzhong Liu Institute of Microbiology, Chinese Academy of Sciences, No 3 Park 1, Beichen West Road, Chaoyang District, Beijing 100101, China Regulo Carlos Llarena Hernandez Facultad de Ciencias Biológicas y Agropecuarias, Peñuela, Universidad Veracruzana, Amatlán de los Reyes, Ver., Mexico Esteve Llop Facultat de Biologia, Universitat de Barcelona, Av. Diagonal, 643, 08028 Barcelona, Spain Csaba Locsmándi Hungarian Natural History Museum, 1087 Budapest, Hungary Deborah Jean Lodge USDA Forest Service, NRS, P.O. Box 1377, Luquillo, Puerto Rico 00773-1377, U.S.A. Michael Loizides P.O. Box 58499, 3734 Limassol, Cyprus László Lőkös Hungarian Natural History Museum, 1087 Budapest, Hungary Jennifer Luangsa-ard National Center for Genetic Engineering and Biotechnology (BIOTEC), NSTDA, 113 Thailand Science Park, Phaholyothin Rd., Klong Nueng, Klong Luang, Pathum Thani 12120, Thailand Matthias Lüderitz AG Geobotanik Schleswig-Holstein & Hamburg, c/o University of Kiel, Olshausenstraße 75, 24098 Kiel, Germany Thorsten Lumbsch Field Museum, 1400 S. Lake Shore Dr., Chicago, Illinois 60605, U.S.A. Matthias Lutz Institute of Evolution and Ecology, University of Tübingen, Auf der Morgenstelle 5, 72076 Tübingen, Germany Dan Mahoney Callaghan Innovation, 69 Gracefield Road, Lower Hutt 5010, New Zealand Ekaterina Malysheva Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Vera Malysheva Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Patinjareveettil Manimohan University of Calicut, Kerala, 673 635, India Yasmina Marin-Felix Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands Guilhermina Marques Centre for the Research and Technology of Agro-Environmental and Biological Sciences, University of Tras-os-Montes and Alto Douro, Laboratory of Mycology and Soil Microbiology, 5000-801 Vila Real, Portugal Rubén Martínez-Gil Ascomycete.org, 36 rue de la Garde, 69005 Lyon, France Guy Marson Musée national d’histoire naturelle, 25 rue Münster, 2160 Luxembourg, Luxembourg Gerardo Mata Instituto de Ecología, A.C., Carretera antigua a Coatepec 351, El Haya, Xalapa 91070, Veracruz, Mexico P. Brandon Matheny University of Tennessee, Knoxville, Tennessee 37996, U.S.A. Geir Harald Mathiassen Tromsø University Museum, University of Tromsø — The Arctic University of Norway, 9037 Tromsø, Norway Neven Matočec Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Helmut Mayrhofer Institute of Biology, University of Graz, Holteiasse 6, 8010 Graz, Austria Mehdi Mehrabi Iranian Research Institute of Plant Protection, Agricultural Research, Education and Extension Organization, Tehran, Iran Ireneia Melo Botanical Garden, National Museum of Natural History and Science, University of Lisbon, Portugal Armin Mešić Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Andrew S. Methven Savannah State University, Savannah, Georgia 31404, U.S.A Otto Miettinen Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Ana M. Millanes Romero Universidad Rey Juan Carlos, 28933 Móstoles, Madrid, Spain Andrew N. Miller Illinois Natural History Survey, University of Illinois Urbana-Champaign, 1816 South Oak Street, Champaign, Illinois 61820-6970, U.S.A. James K. Mitchell Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Roland Moberg Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Pierre-Arthur Moreau Université Lille, Fac. Pharma. Lille, EA4483 IMPECS, 59000 Lille, France Gabriel Moreno Facultad de Biología, Ciencias Ambientales y Química, Universidad de Alcalá, 28805 Alcalá de Henares, Madrid, Spain Olga Morozova Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Asunción Morte Facultad de Biología, Universidad de Murcia, Campus de Espinardo, 30100 Murcia, Spain Lucia Muggia University of Trieste, via L. Giorgieri 10, 34127 Trieste, Italy Guillermo Muñoz González Avda Valvanera N.32, 5D, 26500 Calahorra, La Rioja, Spain Leena Myllys Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland István Nagy Institute of Biology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary László G. Nagy Institute of Biochemistry, BRC-HAS, 6726 Szeged, Temesvari krt. 62, 6726 Szeged, Hungary Maria Alice Neves Universidade Federal de Santa Catarina, Campus Universitário Reitor João David Ferreira Lima, Trindade, Florianópolis, Santa Catarina CEP 88040-900, Brazil Tuomo Niemelä Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Pier Luigi Nimis University of Trieste, via L. Giorgieri 10, 34127 Trieste, Italy Nicolas Niveiro Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Machiel E. Noordeloos Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA, Leiden, The Netherlands Anders Nordin Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Sara Raouia Noumeur Faculty of Natural and Life Sciences, University of Batna 2, 05000 Batna, Algeria Yuri Novozhilov Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Jorinde Nuytinck Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands Esteri Ohenoja Botanical Museum, University of Oulu, Finland Patricia Oliveira Fiuza Universidade Estadual de Feira de Santana, Av. Transnordestina, s/n, Bairro Novo Horizonte, CEP:44036-900, Feira de Santana, Bahia, Brazil Alan Orange National Museum of Wales, Cardiff CF10 3NP, United Kingdom Alexander Ordynets University of Kassel, Heinrich-Plett-Strasse 40, 34132 Kassel, Germany Beatriz Ortiz-Santana USDA Forest Service, Northern Research Station, One Gifford Pinchot Dr, Madison, Wisconsin 53726, U.S.A. Leticia Pacheco Universidad Autónoma Metropolitana-Iztapalapa, Av. San Rafael Atlixco 186, Col. Vicentina, 09340 México, D. F., Mexico Ferenc Pál-Fám Faculty of Agricultural and Environmental Sciences, Kaposvar University, 7400 Kaposvar, Hungary Melissa Palacio Universidade Federal do Rio Grande do Sul, Brazil Zdeněk Palice Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Viktor Papp Szent Istvan University, 1118 Budapest, Menesi st. 44, Hungary Kadri Pärtelv Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Julia Pawlowska Faculty of Biology, Biological and Chemical Research Centre, University of Warsaw, Żwirki i Wigury 101, 02-089 Warsaw, Poland Aurelia Paz Urb. La Llosa, 219, 39509 Villanueva de la Peña, Mazcuerras, Cantabria, Spain Ursula Peintner Institut für Mikrobiologie, Universität Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria Shaun Pennycook Manaaki Whenua Landcare Research, Private Bag 92170, Auckland 1072, New Zealand Olinto Liparini Pereira Universidade Federal de Viçosa, 36570-000, Viçosa, Minas Gerais, Brazil Pablo Pérez Daniëls University of Córdoba, 14071, Córdoba, Spain Miquel À. Pérez-De-Gregorio Capella C/ Pau Casals, 6, 1º, 1ª, 17001, Girona, Spain Carlos Manuel Pérez del Amo C/ Luis de Ulloa, 1, 7º I, 26004 Logroño, Navarra, Spain Sergio Pérez Gorjón Universidad de Salamanca, Avda. Licenciado Mendez Nieto s/n, 37007 Salamanca, Spain Sergio Pérez-Ortega Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Israel Pérez-Vargas Facultad de Farmacia, Universidad de La Laguna, c/ Astrofísico Sánchez s/n 38071 La Laguna, Tenerife, Canary Islands, Spain Brian A. Perry California State University East Bay, Hayward, California 94542, U.S.A. Jens H. Petersen Nøruplundvej 2, 8400 Ebeltoft, Denmark Ronald H. Petersen University of Tennessee, Knoxville, Tennessee 37996, U.S.A. Donald H. Pfister Harvard University, 22 Divinity Avenue, Cambridge MA 02138, U.S.A. Chayanard Phukhamsakda Mae Fah Luang University, Chang Wat Chiang Rai 57100, Thailand Marcin Piątek W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Meike Piepenbring Faculty of Biosciences, Goethe University Frankfurt am Main, Germany Raquel Pino-Bodas Real Jardín Botánico-CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Juan Pablo Pinzón Esquivel Universidad Autónoma de Yucatán, Carretera Mérida-Xmatkuil Km. 15.5, Apdo. Postal: 4-116 Itzimná,C.P: 97100, Merida, Yucatan, Mexico Paul Pirot Rue des Peupliers 10, 6840 Neufchâteau, Belgium Eugene S. Popov Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Orlando Popoff Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina María Prieto Álvaro Universidad Rey Juan Carlos, 28933 Móstoles, Madrid, Spain Christian Printzen Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Abteilung Botanik und Molekulare Evolutionsforschung, Herbarium Senckenbergianum (FR), Senckenberganlage 25, 60325 Frankfurt am Main, Germany Nadezhda Psurtseva Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Witoon Purahong Helmholtz Centre for Environmental Research GmbH — UFZ, Theodor-Lieser-Straße 4, 06120 Halle, Germany Luis Quijada Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. Gerhard Rambold University of Bayreuth, Universitätsstrasse 30, 95447 Bayreuth, Germany Natalia A. Ramírez Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Huzefa Raja University of North Carolina at Greensboro, 435 Sullivan Science Building, PO Box 26170, Greensboro North Carolina 27402-6170, U.S.A. Olivier Raspé Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium Tania Raymundo Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional, Prolongación de Carpio y Plan de Ayala s/n, Miguel Hidalgo, Santo Tomás, 11340 Ciudad de México, Mexico Martina Réblová Institute of Botany, Czech Academy of Sciences, Pruhonice, Czech Republic Yury A. Rebriev Southern Scientific Center, Russian Academy of Sciences, 41 Chehova str., Rostov-on-Don, 344006, Russia Juan de Dios Reyes García Paseo Virgen de Linarejos 6 2 D, Linares, Jaen, Spain Miguel Ángel Ribes Ripoll Avda. Pablo Neruda 120 F, 2°D, 28018 Madrid, Spain Franck Richard CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry Montpellier, EPHE, INSERM, 1919 Route de Mende, 34293 Montpellier Cédex 5, France Mike J. Richardson Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, United Kingdom Víctor J. Rico Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain Gerardo Lucio Robledo Instituto Multidisciplinario de Biología Vegetal, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Córdoba, Casilla de Correo 495, 5000 Córdoba, Argentina Flavia Rodrigues Barbosa Universidade Federal de Mato Grosso, Av. Alexandre Ferronato, 1200, Setor Industrial, Sinop, Mato Grosso, Brazil Cristina Rodriguez-Caycedo UT Southwestern Medical Center, Dallas, Texas 75390, U.S.A. Pamela Rodriguez-Flakus W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Anna Ronikier W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Krakow, Poland Luis Rubio Casas Ul. Andrieja Sacharowa 1/1, 30-806 Kraków, Poland Katerina Rusevska Institute of Biology, Faculty of Natural Science and Mathematics, Ss. Cyril and Methodius University, Arhimedova 5, 1000 Skopje, Republic of Macedonia Günter Saar Dammenmühle 7, 77933 Lahr-Sulz, Germany Irja Saar Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Isabel Salcedo University of the Basque Country (UPV/EHU), Apdo 644, 48080 Bilbao, Spain Sergio M. Salcedo Martínez Facultad de Ciencias Biológicas, UANL. Ave. Pedro de Alba s/n esq. Manuel Barragán Cd. Universitaria, San Nicolás de los Garza Nuevo León, CP. 66451, Mexico Carlos A. Salvador Montoya Instituto de Botánica del Nordeste, Universidad Nacional de Nordeste-Consejo Nacional de Investigaciones Científicas y Técnicas, Sargento Cabral 2131, CC 209, Corrientes Capital, Argentina Santiago Sánchez-Ramírez University of Toronto, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada J. Vladimir Sandoval-Sierra Health Science Centre, University of Tennessee, U.S.A. Sergi Santamaria Facultat de Biociències, Edifici C, Despatx C1/331, Campus de la UAB, 08193 Bellaterra, (Cerdanyola del Vallès), Barcelona, Spain Josiane Santana Monteiro Botany Coordination, Museu Paraense Emílio Goeldi, 66077-830, Belém, Pará, Brazil Hans Josef Schroers Agricultural Institute of Slovenia, Hacquetova ulica 17, 1000 Ljubljana, Slovenia Barbara Schulz Institute of Microbiology, Technische Universität Braunschweig, Germany Geert Schmidt-Stohn Burgstr. 25, 29553 Bienenbüttel, Germany Trond Schumacher University of Oslo, P.O.Box 1066, Blindern, 0316 Oslo, Norway Beatrice Senn-Irlet RU Biodiversity and Conservation Biology, Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Zürcherstr. 111, 8903 Birmensdorf, Switzerland Hana Ševčíková Moravian Museum, Zeny trh 6, 659 37 Brno, Czech Republic Oleg Shchepin Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Takashi Shirouzu Graduate School of Bioresources, Mie University, 1577 Kurima-machiya, Tsu, Mie 514-8507, Japan Anton Shiryaev Institute of Plant & Animal Ecology Ural Branch of the Russian Academy of Sciences, 620144 Ekaterinburg, 8 March str., 202/3, Russia Klaus Siepe Geeste 133, 46342 Velen, Germany Esteban B. Sir Fundación Miguel Lillo, Consejo Nacional de Investigaciones Científicas y Técnicas, Miguel Lillo 251, San Miguel de Tucumán 4000, Tucumán, Argentina Mohammad Sohrabi Iranian Research Organization for Science and Technology, P.O. Box 33535111, Tehran, Iran Karl Soop Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Viacheslav Spirin Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Toby Spribille University of Alberta, Edmonton, Alberta T6G 2R3, Canada Marc Stadler Helmholtz Centre for Infection Research, Inhoffenstrasse 7, 38124 Braunschweig, Germany Joost Stalpers Torenlaan 43, 3742CR Baarn, The Netherlands Soili Stenroos Finnish Museum of Natural History, P.O. Box 7, 00014 University of Helsinki, Finland Ave Suija Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Stellan Sunhede Hökaskog Sandbacken 1, 533 92 Lundsbrunn, Sweden Sten Svantesson University of Gothenburg, Box 461, 40530 Göteborg, Sweden Sigvard Svensson Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Tatyana Yu. Svetasheva (1) Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia; (2) Department of Technologies of Living Systems, Tula State Lev Tolstoy Pedagogical University, Lenin ave. 125, Tula, 300026, Russia Krzysztof Świerkosz Museum of Natural History, Wrocław University, ul. H. Sienkiewicza 5, 50-335 Wrocław, Poland Heidi Tamm Institute of Ecology and Earth Sciences, 40 Lai Street, Tartu 51005, Estonia Hatira Taskin Faculty of Agriculture, University of Çukurova, 01330 Adana, Turkey Adrien Taudière CEFE UMR5175, CNRS, Université de Montpellier, Université Paul-Valéry Montpellier, EPHE, INSERM, 1919 Route de Mende, 34293 Montpellier Cédex 5, France Jan-Olof Tedebrand Sundsvall Mycological Society, Medelpad, Sweden Raúl Tena Lahoz C/Arreñales del Portillo B 21 1°D, 44003 Teruel, Spain Marina Temina Institute of Evolution, University of Haifa, Aba Khoushi Ave. 199, Mt. Carmel, Haifa 3498838, Israel Arne Thell Biological Museum, Lund University, Box 117, 221 00 Lund, Sweden Marco Thines Faculty of Biosciences, Goethe University Frankfurt am Main, Germany Göran Thor Swedish University of Agricultural Sciences, P. O. Box 7044, 750 07 Uppsala, Sweden Holger Thüs State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany Leif Tibell Evolutionary Biology Centre, Norbyvägen 18D, 75236 Uppsala, Sweden Sanja Tibell Evolutionary Biology Centre, Norbyvägen 18D, 75236 Uppsala, Sweden Einar Timdal Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, 0318 Oslo, Norway Zdenko Tkalčec Ruđer Bošković Institute, Bijenička cesta 54, 10000 Zagreb, Croatia Tor Tønsberg University of Bergen, Allégaten 41, P.O. Box 7800, 5020 Bergen, Norway Gérard Trichies 5, impasse des Écoles S7700 Neufchef, France Dagmar Triebel Botanische Staatssammlung München, Menzinger Straße 67, 80638 München, Germany Andrei Tsurykau (1) F. Skorina Gomel State University, Sovetskaja Str. 104, 246019 Gomel, Belarus; (2) Institute of Natural Sciences, Samara National Research University, Moskovskoye shosse 34, 443086 Samara, Russia Rodham E. Tulloss Herbarium Amanitarum Rooseveltensis, P. O. Box 57, Roosevelt, New Jersey 08555-0057, U.S.A. Veera Tuovinen University of Alberta, Edmonton, Alberta T6G 2R3, Canada Miguel Ulloa Sosa Universidad Nacional Autónoma de México, Tercer Circuito/Sin Número, Ciudad Universitaria, Ciudad de México, C. P. 04510, Mexico Carlos Urcelay Instituto Multidisciplinario de Biología Vegetal, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Córdoba, Casilla de Correo 495, 5000 Córdoba, Argentina François Valade 11 rue haras, le boqueteau, 91240 Saint-Michel-sur-Orge, France Ricardo Valenzuela Garza Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional, Prolongación de Carpio y Plan de Ayala s/n, Miguel Hidalgo, Santo Tomás, 11340 Ciudad de México, Mexico Pieter van den Boom Arafura 16, 5691 JA Son, The Netherlands Nicolas Van Vooren Ascomycete.org, 36 rue de la Garde, 69005 Lyon, France Aida M. Vasco-Palacios Escuela de Microbiología, Universidad de Antioquia, AA1226, Fundación Biodiversa Colombia, Medellín, Colombia Jukka Vauras Herbarium TUR, Biodiversity Unit, University of Turku, 20014 Turku, Finland Juan Manuel Velasco Santos 3 C/Pontevedra, 18, 1º C, 37003 Salamanca, Spain Else Vellinga 861 Keeler Avenue, Berkeley, California 94708, U.S.A. Annemieke Verbeken Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium Per Vetlesen Norges Sopp- og Nyttevekstforbund, Schweigaards gate 34F, 0191 Oslo, Norway Alfredo Vizzini University of Torino, Viale P.A. Mattioli 25, 10125, Torino, Italy Hermann Voglmayr University of Vienna, Rennweg 14, 1030 Vienna, Austria Sergey Volobuev Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia Wolfgang von Brackel Kirchenweg 2, 91341 Röttenbach, Germany Elena Voronina Lomonosov Moscow State University, Biology Faculty, Moscow, Russia Grit Walther Arvid-Harnack-Str. 4, 07743 Jena, Germany Roy Watling Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, United Kingdom Evi Weber Blaihofstr. 42, 72074 Tübingen, Germany Mats Wedin Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Øyvind Weholt Nord University, Nesna, 8700 Nesna, Norway Martin Westberg Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden Eugene Yurchenko Polessky State University, Dnyaprouskai flatylii str. 23, 225710, Pinsk, Belarus Petr Zehnálek Faculty of Science, Charles University, Benátská 2, 128 01 Praha 2, Czech Republic Huang Zhang Kunming University of Science & Technology, Kunming, China Mikhail P. Zhurbenko Komarov Botanical Institute of the Russian Academy of Sciences, 2 Prof. Popov Street, St. Petersburg, 197376, Russia The following institutions are also supporting the present text (institutional support means that committees from the mentioned) Asociación Micológica Carlos Spegazzini (Argentina) Austrian Mycological Society (Austria) Croatian Mycological Society (Croatia) Committee of the Czech Scientific Society for Mycology (Czech Republic) The Netherlands Mycological Society (The Netherlands) Sociedad Española de Liquenología (Portugal/Spain) Iberian Mycological Society (which is under constitution, Portugal/Spain) Federación de Asociaciones Micológicas Andaluzas (FAMA) (Spain) Asociación Botánica y Micológica de Jaén (Spain) Asociación Micológica Hispalense Muscaria (Spain) Societat Micològica Valenciana (Spain)

Rinodinais a widespread, polyphyletic genus of crustosePhysciaceaewithc. 300 species worldwide. A major missing link in understanding its global biogeography has been eastern Asia where the genus has never been systematically revised. Here we review specimen and literature records forRinodinafor north-eastern Asia (Russian Far East, Japan and the Korean Peninsula) and recognize 43 species. We describe two species,R. hypobadiaandR. orientalis, as new to science.Rinodina hypobadiais distinguished by its pigmented hypothecium,Dirinaria-type ascospores and pannarin in both thallus and epihymenium.Rinodina orientalisis characterized by its erumpent apothecia that remain broadly attached, with discs sometimes becoming convex and excluding the thalline margins, ascospores belonging to thePhyscia-type and secondary metabolites absent. Nine other species are reported from the region for the first time. These includeR. dolichospora,R. freyi,R. metaboliza,R. sicula,R. subminutaandR. willeyi. Of particular biogeographical interest are three additional new records that have western North American–eastern Asian distributions: the corticolous speciesR. endospora,R. macrosporaandR. megistospora. Six species have the better known eastern North American–eastern Asian distributions:R. ascociscana(syn.R. akagiensis,R. melancholica),R. buckii,R. chrysidiata,R. subminuta,R. tenuis(syn.R. adirondackii) andR. willeyi, and two have eastern North American–eastern Asian–European distributions:R. excrescensandR. moziana(syn.R. destituta,R. vezdae). Our study begins to close one of the largest gaps in our knowledge of circumboreal species distributions inRinodinaand, together with previous studies in North America and Europe, provides new insights into circumboreal crustose lichen biogeography.Rinodina cinereovirens(syn.R. turfaceavar. cinereovirens) is also reported as new to North America.

Stable, long-term interactions between fungi and algae or cyanobacteria, collectively known as lichens, have repeatedly evolved complex architectures with little resemblance to their component parts. Lacking any central scaffold, the shapes they assume are casts of secreted polymers that cement cells into place, determine the angle of phototropic exposure and regulate water relations. A growing body of evidence suggests that many lichen extracellular polymer matrices harbor unicellular, non-photosynthesizing organisms (UNPOs) not traditionally recognized as lichen symbionts. Understanding organismal input and uptake in this layer is key to interpreting the role UNPOs play in lichen biology. Here, we review both polysaccharide composition determined from whole, pulverized lichens and UNPOs reported from lichens to date. Most reported polysaccharides are thought to be structural cell wall components. The composition of the extracellular matrix is not definitively known. Several lines of evidence suggest some acidic polysaccharides have evaded detection in routine analysis of neutral sugars and may be involved in the extracellular matrix. UNPOs reported from lichens include diverse bacteria and yeasts for which secreted polysaccharides play important biological roles. We conclude by proposing testable hypotheses on the role that symbiont give-and-take in this layer could play in determining or modifying lichen symbiotic outcomes., Polysaccharide composition of lichens is reviewed with an emphasis on extracellular matrix and potential input and uptake by organismal components of the matrix.

Holien, H., Palice, Z., Bjork, C. R., Goward, T. & Spribille, T. 2016. Lecidea coriacea sp. nov., a lichen species from oldgrowth boreal and montane forests in Europe and North America. — Herzogia 29: 412–420. Lecidea coriacea is described as new to science from the boreal forests of Europe and montane conifer forests of northwestern North America. It is probably related to some of the species currently assigned to the genus Puttea, but is included in Lecidea awaiting a more thorough revision of this group. The species is characterized by pale to dark brown apothecia, plusiosporic asci and by the production of secalonic acid A in the hypothecium causing a golden yellow reaction with KOH. Lecidea coriacea seems to be a species of oldgrowth forests and is threatened by forestry. It often grows on old trees of Betula, Picea and Salix or on old conifer snags. Notes on similar species and other plusiosporic epiphytic and lignicolous species in boreal forests are given.

Grube, M., Obermayer, W., Mayrhofer, H. & Spribille, T. 2016. Josef Hafellner — a life amongst lichens and their parasites. — Herzogia 29: 213–234. The impact of Josef Hafellners scientific work on the systematics of lichenized ascomycetes and on the knowledge on lichenicolous fungi is discussed and the value of his extensive collections (c. 100,000 lichens including c. 8,000 lichenicolous fungi) is highlighted. A bibliography (1975–2016) is presented.

Since the late 1800s, mycologists have been detecting fungi above and beyond the assumed single fungus in lichen thalli [1-6]. Over the last century, these fungi have been accorded roles ranging from commensalists to pathogens. Recently, Cyphobasidiales yeasts were shown to be ubiquitous in the cortex layer of many macrolichens [7], but for most species, little is known of their cellular distribution and constancy beyond visible fruiting structures. Here, we demonstrate the occurrence of an additional and distantly related basidiomycete, Tremella, in 95% of studied thalli in a global sample of one of the most intensively studied groups of lichens, the wolf lichens (genus Letharia). Tremella species are reported from a wide range of lichen genera [8], but until now, their biology was deduced from fruiting bodies (basidiomata) formed on lichen thalli. Based on this, they have been thought to be uncommon to rare, to occur exclusively in a hyphal form, and to be parasitic on the dominant fungal partner [9, 10]. We show that, in wolf lichens, Tremella occurs as yeast cells also in thalli that lack basidiomata and infer that this is its dominant stage in nature. We further show that the hyphal stage, when present in Letharia, is in close contact with algal cells, challenging the assumption that lichen-associated Tremella species are uniformly mycoparasites. Our results suggest that extent of occurrence and cellular interactions of known fungi within lichens have historically been underestimated and raise new questions about their function in specific lichen symbioses.

Lichens exhibit varying degrees of specialization with regard to the surfaces they colonize, ranging from substrate generalists to strict substrate specialists. Though long recognized, the causes and consequences of substrate specialization are poorly known. Using a phylogeny of a 150–200 Mya clade of lichen fungi, we asked whether substrate niche is phylogenetically conserved, which substrates are ancestral, whether specialists arise from generalists or vice versa and how specialization affects speciation/extinction processes. We found strong phylogenetic signal for niche conservatism. Specialists evolved into generalists and back again, but transitions from generalism to specialism were more common than the reverse. Our models suggest that for this group of fungi, ‘escape’ from specialization for soil, rock and bark occurred, but specialization for wood foreclosed evolution away from that substrate type. In parallel, speciation models showed positive diversification rates for soil and rock dwellers but not other specialists. Patterns in the studied group suggest that fungal substrate specificity is a key determinant of evolutionary trajectory for the entire lichen symbiosis.

Contributing to the process of reassigning lecideoid lichens to natural taxa, we assessed phylogenetic relationships and species delimitation in the Calvitimela aglaea complex (Tephromelataceae) using DNA sequence data and morphological/anatomical and chemical characters. Phylogenetic analysis of nuclear (ITS, MCM7, TEF1-α) and mitochondrial (ribosomal SSU) DNA sequences revealed Mycoblastus as sister to a strongly supported clade comprising Calvitimela, Tephrolema and Violella. Species of these three genera fall into six strongly supported subclades with low backbone resolution. Two of these are represented by Tephromela and Violella, which are readily circumscribed morphologically. The remaining four subclades encompass lineages that have until now been assigned to Calvitimela. While Tephromela and Violella as currently circumscribed are recovered as monophyletic in our analyses, Calvitimela is paraphyletic, with four deeply divergent clades. We recognize these four clades as subgenera Calomela, Calvitimela, Paramela and Severidea. Our molecular results further support the recognition of two recently discovered sterile crusts as new species, Calvitimela cuprea and C. livida, distinguished from previously known species by their production of asexual diaspores and from each other by secondary metabolite chemistry. We also report Calvitimela perlata as new for continental North America.

A new, highly effective primer pair to exclude algae when amplifying nuclear large ribosomal subunit (LS) DNA from lichens

The term symbiosis was first used in biology to describe the ‘living together’ of fungi and algae in lichens. For much of the 20th century, the fungal partner was assumed to be invested with the ability to produce the lichen body plan in presence of a photosynthesizing partner. However, studies of fungal evolution have uncovered discordance between lichen symbiotic outcomes and genome evolution of the fungus. At the same time, evidence has emerged that the structurally important lichen cortex contains lichen-specific, single-celled microbes, suggesting it may function like a biofilm. Together, these observations suggest we may not have a complete overview of symbiotic interactions in lichens. Understanding phenotype development and evolution in lichens will require greater insight into fungal–fungal and fungal–bacterial interplay and the physical properties of the cortex.

Spilonema was originally described to accommodate an unusual group of cyanolichens with thread-like, cushion-forming thalli, and has long been placed in Coccocarpiaceae based on ascomatal development. However, Spilonema is the only genus of Peltigerales to include species lichenized with the cyanobacterial genus Stigonema, and the evolutionary relationships of Spilonema to other genera in the family have yet to be tested using molecular data. We present evidence from combined nuclear 28S, 18S and mitochondrial 12S rDNA to confirm the placement of the core species of Spilonema (S. paradoxum and S. revertens) in Coccocarpiaceae. Our data further show that despite possessing a different genus of photobiont (Scytonema), the north Pacific endemic genus Spilonemella must be included within Spilonema, suggesting that closely related species of the genus have changed photobionts in the course of evolution. However, we recovered Spilonema dendroides, one of the only lichens known to associate with the cyanobacterial genus Hyphomorpha, as only distantly related to the Coccocarpiaceae. The evolutionary relationships of this species are as yet unclear but it may occupy a basal position in the Peltigerales. We create for this species the new genus Erinacellus T. Sprib., Muggia & Tønsberg.

Question The mountains of NW North America are home to natural meadows that cover large areas at the timberline. The vegetation of these meadows has been the topic of few local studies, and little is known of species composition turnover across mountain ranges and along the latitudinal gradient. We sampled and analyzed community composition along a 1000-km gradient to answer the following questions: (1) what are the main community types in timberline meadows; (2) what are the most important abiotic gradients underlying their composition; and (3) how is species diversity related to environmental factors and how does it change along the latitudinal gradient? Location SE and central British Columbia (Canada), NW Montana (USA). Methods We recorded data on cover and presence/absence of vascular plant, bryophyte and lichen species, as well as structure and environmental properties in 4 m × 4 m plots (n = 112). Using the OptimClass approach, we classified plots into distinct community types. We used NMDS and multiple regressions to analyse the relationship between composition, species richness and environmental and geographic gradients. Results Despite drawing from a large geographic area, our classification revealed only four meadow communities. Mesic meadows were the most frequent and had a pronounced species turnover along latitude, resulting in two distinct community types. In addition, a three-dimensional ordination showed that vegetation structural properties, macroclimate and geographic predictors were important underlying gradients for species composition. Vascular species richness increased with soil pH, and exhibited a quadratic relationship with litter cover and latitude. Furthermore, in the latter case, it increased towards the southern and northern ends of our gradient. Conclusions Detection of only four meadow communities along the 1000-km sampling gradient suggests a relatively recently established flora and a lack of regional differentiation consistent with the low level of endemism in the region. Higher vascular species diversity at the northern and southern edges of our gradient may reflect the geographic proximity to areas unglaciated during the Pleistocene, and suggests that areas farthest from the unglaciated edge may be experiencing colonization debt.

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.

Lichens assemble in three parts Lichen growth forms cannot be recapitulated in the laboratory by culturing the plant and fungal partners together. Spribille et al. have discovered that the classical binary view of lichens is too simple. Instead, North American beard-like lichens are constituted of not two but three symbiotic partners: an ascomycetous fungus, a photosynthetic alga, and, unexpectedly, a basidiomycetous yeast. The yeast cells form the characteristic cortex of the lichen thallus and may be important for its shape. The yeasts are ubiquitous and essential partners for most lichens and not the result of lichens being colonized or parasitized by other organisms. Science , this issue p. 488

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 ...

Sheard, J. W., Lendemer, J. C., Spribille, T., Thor, G. & Tonsberg, T. 2012. Further contributions to the genus Rinodina (Physciaceae, Lecanoromycetidae): two species new to science and a new record for the Canadian High Arctic. — Herzogia 25: 125–143. Two species of Rinodina are described as new to science (R. buckii and R. bullata) and a third is reported for the first time from arctic Canada (R. ochracea). Rinodina buckii, a sorediate species that resembles R. willeyi is described from disjunct localities in East Asia and the Appalachian Mountains of eastern North America. Rinodina bullata, a species with a peculiar minutely areolate thallus that is frequently sterile, is described from the central Appalachian Mountains of eastern North America. Rinodina ochracea, originally described from Greenland, is reported from the Northwest Territories in Canada. The newly described species are illustrated with both colour light micrographs (CLMs) and scanning electron micrographs (SEMs), and maps of t...

The Hypogymnia austerodes group is an assemblage of mostly boreal and arctic-alpine lichens traditionally comprising three wide-ranging sorediate and/or isidiate species, H. austerodes, H. bitteri and H. subobscura. Here we describe four additional members of this group – H. dichroma sp. nov., H. protea sp. nov., H. salsa sp. nov. and H. verruculosa sp. nov. – which have until now been treated within H. austerodes s. lat. Our new species are primarily epiphytic, and are currently known only from Cordilleran western North America. They differ from other members of the H. austerodes group both morphologically and in their virtually consistent production of apinnatic acid. Hypogymnia protea is by far the most variable of the new species, uniting two putative taxa here designated as morphs “disjuncta” and “tessellata”. Reliable discrimination of the segregate species requires careful attention to their vegetative reproductive structures, which are described in detail. A key to all treated species and...

The crustose lichen genusMycoblastusin the Northern Hemisphere includes eight recognized species sharing large, simple ascospores produced 1–2 per ascus in strongly pigmented biatorine apothecia. The monophyly ofMycoblastusand the relationship of its various species toTephromelataceaehave never been studied in detail. Data from ITS rDNA and the genes coding for translation elongation factor 1-α and DNA replication licensing factor mini-chromosome maintenance complex 7 support the distinctness ofMycoblastuss. str. from the core of theTephromelataceae, but recoverM. fucatusand an undescribed Asian species as strongly supported within the latter group. We propose accommodating these two species in a new genus,Violella, which is characterized by its brownish inner ascospore walls, Fucatus-violet hymenial pigment granules and secondary chemistry, and discuss the position ofViolellarelative toCalvitimelaandTephromela. We describe the new speciesViolella wangiiT. Sprib. & Goffinet to accommodate a new species with roccellic acid from Bhutan, China, India and the Russian Far East. We also excludeMycoblastus indicusAwasthi & Agarwal from the genusMycoblastusand propose for it the new combinationMalmidea indica(Awasthi & Agarwal) Hafellner & T. Sprib.

The new lichenicolous fungus Llimoniella cinnabarinae growing on Ramboldia cinnabarina from Alaska is described. The lack of excipular or epihymenial K+ purplish, violet or green pigmentsplaces it near the recently described L. phaeophysciae group. Its affinity with Llimoniella and othermembers of lichenicolous Helotiales is discussed. Key words: lichenicolous fungus, lichens, North America, taxonomy Introduction Thegenus Llimoniella wasintroducedfortwolichenicolous fungi growing on terricolouslichens of gypsum soils (Hafellner N Diederich et al .2010). The original concept of Llimoniella hasrecentlybeenenlargedtoincludeagroupof taxa, referred to as the L. phaeophysciae group, which does not exhibit the K+ violet,purplish or green pigments in the epi-hymenium and excipulum but are other-wise typical of Llimoniella (Diederich et al .2010). During the course of recent work onlichens and lichenicolous fungi in Alaska(e.g., Spribille

Helmut Mayrhofer has been contributing to understanding the systematics and diversity of lichenized fungi for over four decades. In this paper introducing a Festschrift on the occasion of his 65th birthday, we review the impact of Helmut Mayrhofer's scientific work on the systematics of lichenized fungi with special emphasis on the family Physciaceae and highlight his studies of the lichen flora of the Balkan Peninsula, the Alps and other regions. We also present a bibliography of his publications from 1973 to the present.

The lichen flora of southeast Alaska has been explored for over 100 years, but remains poorly known. In the first survey of its kind from the region, we report 766 taxa of lichens and lichenicolous fungi from the Klondike Gold Rush National Historic Park. Coming from a park only 53 km 2 in size, this represents one of the largest numbers of lichenized and lichenicolous fungi per unit area ever reported and the largest number ever reported from any United States National Park of any size. One lichen genus, four lichen species and one species of lichenicolous fungus are described as new to science: Steineropsis alaskana gen. et sp. nov. (Peltigerales), Coccotrema hahriae and Pertusaria mccroryae (both Pertusariales), Stereocaulon klondikense (Lecanorales) and Corticifraga scrobiculatae (Lecanoromycetes incertae sedis). The new combinations Coccotrema minutum and Nesolechia fusca are made and Parmelia elongata is reduced to synonymy under Hypogymnia duplicata. Seventy-five taxa could not be assigned a species name at this time and represent a pool of critical and/or potentially undescribed taxa. Thirty-four taxa are new or confirmed for North America, including seventeen lichen taxa, one species of saprophytic Dothideomycetes, and sixteen species of lichenicolous fungi. Five taxa of eastern North American distribution are reported for the west for the first time. A total of 196 taxa are new for Alaska. We report the presence of novel secondary chemical compounds in Bryoria, Cladonia, Hypogymnia and Pertusaria. Based on a Chao analysis of single and two-time occurrences we estimate we have captured not more than 83% of the macrolichen and 64% of the microlichen flora, with the total flora likely exceeding 1000 taxa. We provide an overview of the lichen inventories with highest species number worldwide and discuss the Klondike in the context of broader patterns in lichen richness. Global models of climate change in the coming century are unanimous in predicting greater temperature increases at higher latitudes than at low latitudes. We hope that our data will lead to a reconsideration of the potential extent of biodiversity at high latitudes.

Elixia cretica T. Sprib. & Lumbsch is described as a new species from the mountains of western Crete. The second member of the previously monotypic genus and only the third member of the family Elixiaceae, it is distinguished by its surficial thallus, larger ascospores and corticolous habit. Molecular phylogenetic analysis based on a sequence of mitochondrial small subunit DNA confirms the position of the new species. We also report E. flexella from New Hampshire (USA) as new to eastern North America.

Hypogymnia metaphysodes was first described from Japan and Sakhalin, and later reported from western North America. Here we show that the North American material currently referred to H. metaphysodes differs from that species not only morphologically and chemically, but also in ascospore size and shape. We also show that the North American material is in fact heterogeneous, and can be assigned to two well-defined species here described as new: Hypogymnia recurva sp. nov. and H. wilfiana sp. nov. Both of these lichens contain distinctive secondary metabolites: vittatolic acid in the case of H. recurva, and 2-methylene-3-carboxy-18-hydroxynonadecanoic acid (“apinnatic acid”), reported here for the first time from Hypogymnia , in H. wilfiana. Both of our new species are so far known only from western North America, where they occur primarily as epiphytes on the branches of conifers. Hypogymnia metaphysodes s. str. has not yet been reliably reported from this region and should be excluded from the North American lichen flora.

The phylogeny of the Lecanora varia group is inferred from ITS sequences using Bayesian and ML phylogenetic analysis methods. According to our data, usnic acid-containing, corticolous Lecanora species do not form a monophyletic group but occur in three major clades together with other groups of Lecanora and Protoparmeliopsis. The new combination Lecanora filamentosa (Stirt.) Elix & Palice is proposed. The new species Lecanora schizochromatica is described from northwestern North America. It produces atranorin as a major substance and is closely related to the morphologically and chemically similar L. filamentosa. The American Biatora pullula Tuck. is synonymised with Lecanora anopta Nyl., which is hereby reported for continental North America for the first time. The phylogenetic relationships between the major clades of Lecanora are still largely unresolved and require more intensive taxon and character sampling. peerReviewed