Lichens and Associated Fungi from Glacier Bay National Park, Alaska
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The Lichenologist (2020), 52,61–181 doi:10.1017/S0024282920000079 Standard Paper Lichens and associated fungi from Glacier Bay National Park, Alaska Toby Spribille1,2,3 , Alan M. Fryday4 , Sergio Pérez-Ortega5 , Måns Svensson6, Tor Tønsberg7, Stefan Ekman6 , Håkon Holien8,9, Philipp Resl10 , Kevin Schneider11, Edith Stabentheiner2, Holger Thüs12,13 , Jan Vondrák14,15 and Lewis Sharman16 1Department of Biological Sciences, CW405, University of Alberta, Edmonton, Alberta T6G 2R3, Canada; 2Department of Plant Sciences, Institute of Biology, University of Graz, NAWI Graz, Holteigasse 6, 8010 Graz, Austria; 3Division of Biological Sciences, University of Montana, 32 Campus Drive, Missoula, Montana 59812, USA; 4Herbarium, Department of Plant Biology, Michigan State University, East Lansing, Michigan 48824, USA; 5Real Jardín Botánico (CSIC), Departamento de Micología, Calle Claudio Moyano 1, E-28014 Madrid, Spain; 6Museum of Evolution, Uppsala University, Norbyvägen 16, SE-75236 Uppsala, Sweden; 7Department of Natural History, University Museum of Bergen Allégt. 41, P.O. Box 7800, N-5020 Bergen, Norway; 8Faculty of Bioscience and Aquaculture, Nord University, Box 2501, NO-7729 Steinkjer, Norway; 9NTNU University Museum, Norwegian University of Science and Technology, NO-7491 Trondheim, Norway; 10Faculty of Biology, Department I, Systematic Botany and Mycology, University of Munich (LMU), Menzinger Straße 67, 80638 München, Germany; 11Institute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK; 12Botany Department, State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany; 13Natural History Museum, Cromwell Road, London SW7 5BD, UK; 14Institute of Botany of the Czech Academy of Sciences, Zámek 1, 252 43 Průhonice, Czech Republic; 15Department of Botany, Faculty of Science, University of South Bohemia, Branišovská 1760, CZ-370 05 České Budějovice, Czech Republic and 16Glacier Bay National Park & Preserve, P.O. Box 140, Gustavus, Alaska 99826, USA Abstract Lichens are widely acknowledged to be a key component of high latitude ecosystems. However, the time investment needed for full inventories and the lack of taxonomic identification resources for crustose lichen and lichenicolous fungal diversity have hampered efforts to fully gauge the depth of species richness in these ecosystems. Using a combination of classical field inventory and extensive deployment of chemical and molecular analysis, we assessed the diversity of lichens and associated fungi in Glacier Bay National Park, Alaska (USA), a mixed landscape of coastal boreal rainforest and early successional low elevation habitats deglaciated after the Little Ice Age. We collected nearly 5000 specimens and found a total of 947 taxa, including 831 taxa of lichen-forming and 96 taxa of licheni- colous fungi together with 20 taxa of saprotrophic fungi typically included in lichen studies. A total of 98 species (10.3% of those detected) could not be assigned to known species and of those, two genera and 27 species are described here as new to science: Atrophysma cyanomelanos gen. et sp. nov., Bacidina circumpulla, Biatora marmorea, Carneothele sphagnicola gen. et sp. nov., Cirrenalia lichenicola, Corticifraga nephromatis, Fuscidea muskeg, Fuscopannaria dillmaniae, Halecania athallina, Hydropunctaria alas- kana, Lambiella aliphatica, Lecania hydrophobica, Lecanora viridipruinosa, Lecidea griseomarginata, L. streveleri, Miriquidica gyrizans, Niesslia peltigerae, Ochrolechia cooperi, Placynthium glaciale, Porpidia seakensis, Rhizocarpon haidense, Sagiolechia phaeospora, Sclerococcum fissurinae, Spilonema maritimum, Thelocarpon immersum, Toensbergia blastidiata and Xenonectriella nephromatis.An additional 71 ‘known unknown’ species are cursorily described. Four new combinations are made: Lepra subvelata (G. K. Merr.) T. Sprib., Ochrolechia minuta (Degel.) T. Sprib., Steineropsis laceratula (Hue) T. Sprib. & Ekman and Toensbergia geminipara (Th. Fr.) T. Sprib. & Resl. Thirty-eight taxa are new to North America and 93 additional taxa new to Alaska. We use four to eight DNA loci to validate the placement of ten of the new species in the orders Baeomycetales, Ostropales, Lecanorales, Peltigerales, Pertusariales and the broader class Lecanoromycetes with maximum likelihood analyses. We present a total of 280 new fungal DNA sequences. The lichen inventory from Glacier Bay National Park represents the second largest number of lichens and associated fungi documented from an area of comparable size and the largest to date in North America. Coming from almost 60°N, these results again underline the potential for high lichen diversity in high latitude ecosystems. Key words: biodiversity, evolution, floristics, key, latitudinal diversity gradient, molecular, new species, phylogenetics, symbiosis, taxonomy, temperate rainforest (Accepted 8 October 2019) Author for correspondence: Toby Spribille. E-mail: [email protected] Cite this article: Spribille T et al. (2020) Lichens and associated fungi from Glacier Bay National Park, Alaska. Lichenologist 52,61–181. https://doi.org/10.1017/ S0024282920000079 © British Lichen Society 2020. This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited. Downloaded from https://www.cambridge.org/core. IP address: 73.181.224.165, on 25 May 2020 at 14:45:03, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282920000079 62 Toby Spribille et al. Table of Contents Introduction Introduction 62 The landscapes of south-east Alaska are best known for their most The present study 63 striking macrofeatures: snow-capped mountains, misty saltwater Materials and Methods 63 fjords and dark coniferous rainforests. Closer examination reveals Study area 63 that the texture of nearly every terrestrial feature in south-east Climate 63 Alaska is, in one way or another, determined at a much smaller Glaciation and vegetation history 64 scale. Zooming from the landscape view into the canopies of Stratification of study area into target sampling units 65 the coastal rainforests and the tapestry of their outcrops and boul- Specimen analysis 65 der fields reveals a Russian doll of nested ecosystems, one within Molecular data 65 another, within another. At the scale of an ecosystem a human can Phylogenetic trees 66 hold in her hand, it is fungi and bryophytes that form the building Species delimitation and nomenclature 76 blocks of the multicellular canopy, supporting yet another set of Presentation of species data 77 nested dolls of microbial and invertebrate life in their peaks and Comparison between sectors and national parks 77 ravines. It is at this scale, where fungi, algae and bacterial biofilms Results and Discussion 77 meet in a permanently wet, cold milieu, that the south-east Alaskan Comparison of sectors within GLBA 77 temperate rainforest exhibits peak biodiversity. ’ Lichen diversity in the national parks of the greater Lichens, s éixwani to the Tlingit (Edwards 2009), the indigen- Gulf of Alaska region 78 ous people of south-east Alaska, played a role in traditional food Phylogenetic trees 79 and garment dyeing for the residents of these fjords for thousands Descriptions of New Genera and Species 85 of years. In Glacier Bay, the subject of the present paper, lichens Atrophysma T. Sprib. 85 are featured in place names and play an outsized role in the recent Atrophysma cyanomelanos T. Sprib. 85 vegetation history. When the first European collections of lichens Bacidina circumpulla S. Ekman 88 were made here, in the framework of the Harriman Expedition Biatora marmorea T. Sprib. 89 (Cummings 1904), Glacier Bay had only recently undergone a Carneothele Fryday, T. Sprib. & M. Svenss. 91 massive glacial retreat of over 80 km as a result of saltwater glacial Carneothele sphagnicola Fryday, M. Svenss. & Holien 91 erosion. Only a few years later, the American ecologist William Cirrenalia lichenicola Pérez-Ort. 93 Skinner Cooper arrived in Glacier Bay and began a series of stud- Corticifraga nephromatis Pérez-Ort. 93 ies that shaped the textbook description of plant succession Fuscidea muskeg Tønsberg & M. Zahradn. 94 (Cooper 1923), now the longest-running primary succession Fuscopannaria dillmaniae T. Sprib. 96 plot series in the world (Buma et al. 2017). Despite its fame in Halecania athallina Fryday 98 plant ecology, Glacier Bay was neglected by lichen researchers Hydropunctaria alaskana Thüs & Pérez-Ort. 99 in the 20th century. Far fewer collectors have worked here com- Lambiella aliphatica T. Sprib. & Resl 101 pared to other localities in Alaska, for example, the Juneau region Lecania hydrophobica T. Sprib. & Fryday 103 (Krog 1968), Sitka or the north end of the Lynn Canal (see e.g. Lecanora viridipruinosa M. Svenss. & T. Sprib. 105 Spribille et al. 2010). Between the 1899 Harriman Expedition Lecidea griseomarginata Fryday 106 and the beginning of the present study, we could reconstruct Lecidea streveleri T. Sprib. 107 the activity of 17 different collectors or groups of collectors, Miriquidica gyrizans Fryday 110 based on specimens in US, Canadian and Swedish herbaria Niesslia peltigerae Pérez-Ort. 112 (Supplementary Material Table S1, available online). Most