Teuvo Ahti

Epigeic lichen communities of taiga and tundra regions

The major physiognomic and ecological categories of the lichen-rich, epigeic communities in the boreal (taiga) and arctic (tundra) zones are defined and their syntaxonomy and ecology in Europe, Asia and North America is reviewed. In the boreal and hemiarctic areas open, dry, acidophytic lichen woodlands are widespread, especially on sandy habitats. Their epigeic lichen synusiae are usually dominated by four fruticoseCladina species, being extremely homogeneous in species composition and structure throughout the boreal zone, while the dominant trees and the other vascular plant flora of the woodlands are geographically more variable. No phytosociological classification system exists that would cover most of these communities over the circumpolar regions. Very similar communities, though much more poorly known, are found on thin soils over Precambrian rock outcrops. Other sites to produce epigeic lichen communities include open sand dunes, treeless heathlands, drier bogs and many seral stages, like those on road banks. Boreal lichen-rich communities on eutrophic soils may be developed in semiarid regions, in particular. In the Arctic, lichens are common in most communities, and the driest ones are regularly lichen-dominated, whether acidophytic or eutrophytic, chionophytic or achionophytic. Detailed syntaxonomic systems for their classification have been developed, especially in Greenland and Scandinavian mountains (in oroarctic zones in the latter regions). The richest fruticose lichen areas are in continental, hemiarctic timberline regions in northern Siberia and Canada. The southern and middle arctic subzones are also characterized by many macrolichens, such asCetraria cucullata, C. nivalis, Alectoria ochroleuca, andThamnolia vermicularis, but everywhere also small, crustose lichens are common on soil, such asRinodina turfacea andLopadium pezizoideum, which are often overlooked in vegetation analyses. The presence of microlichens and the formation of mosaic micropatterns of soil lichen communities is particularly typical of the northern arctic subzone. The conservation problems of the boreal and arctic lichen communities include overgrazing by reindeer or caribou, which has caused delichenization in some regions, extensive forest and tundra fires, use of heavy transport vehicles in forestry and tundra operations, and, locally, heavy industrial air pollution.

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

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

Molecular data show that Bryoria fremontii and B. tortuosa (Parmeliaceae) are conspecific

Bryoria fremontii and B. tortuosa are the only species in the lichenized ascomycete genus Bryoria known to contain the pulvinic acid derivative vulpinic acid. In B. fremontii this yellow pigment is restricted to the soralia and apothecia, while in B. tortuosa it can occur throughout the thallus. The actual amount of vulpinic acid produced by B. tortuosa is rather variable, however, with intermediate specimens bearing both white and yellow pseudocyphellae. We studied the relationship between the two species with parsimony analysis using four DNA regions: 1) the internal transcribed spacers of the nuclear rDNA including the 5.8S region (ITS), 2) partial sequences from the intergenic spacer of the nuclear rDNA (IGS), 3) partial sequences from the small subunit of the mitochondrial rDNA (mtSSU), and 4) partial sequences from the protein-coding glyceraldehyde-3-phosphate dehydrogenase gene (GAPDH). Our phylogenetic analysis revealed that B. fremontii and B. tortuosa must be regarded as conspecific, but allowing for some genetic differentiation between European and North American populations. Bryoria tortuosa is therefore synonymized with B. fremontii .

New and Interesting Records of Cladonia and their Lichenicolous Fungi from the Andean Cloud Forest in Bolivia

The paper deals with 18 species of the lichen genus Cladonia, including eight species new to Bolivia: C. cf. borbonica, C. coccifera, C. confragosa, C. dactylota, C. granulosa, C. kriegeri, C. merochlorophaea, and C. squamosa. Furthermore, C. kriegeri is recorded here from its second and third localities in the world. Also five lichenicolous fungi are reported; among them Arthrorhaphis aeruginosa, Polycoccum microcarpum and Sphaerellothecium cladoniae are new to the southern hemisphere, and Roselliniella cladoniae and Tremella cladoniae are new to Bolivia. Additionally, Polycoccum microcarpum is reported for the first time from Slovakia and Ukraine. Cladonia confragosa is reported as a new host for Sphaerellothecium cladoniae and Tremella cladoniae.

Convergent evolution in Cladonia gracilis and allies

Members of the Cladonia gracilis group of lichen fungi are common terrestrial lichens where morphological features are more similar between members of the C. gracilis species complex and allied species outside the complex than they are between subspecies within the complex. The objectives of this study were to examine whether the Cladonia gracilis species complex is monophyletic, to determine whether morphological similarity is supported by genetic variation, and to examine the utility of the polyketide synthase (PKS) gene for phylogenetic studies among closely related species. Two loci, the ketosynthase region of the PKS gene and the internal transcribed spacer (ITS) region of nuclear ribosomal DNA, were sequenced and analysed by Maximum Parsimony, Bayesian and haplotype network analyses. Functional differences were also inferred through ITS2 RNA secondary structures and non-synonymous changes in translated PKS amino acid sequences. The monophyly of the C. gracilis complex is supported by 71% bootstrap in the ITS phylogeny, and 92% bootstrap with greater than 95% posterior probability in the PKS phylogeny. Morphological similarity is not always supported by genetic similarity. The PKS gene is less variable than the ITS but the PKS supports species hypotheses that are reflected in the ITS2 RNA model. We conclude that monophyly of the C. gracilis complex can be supported if C. cornuta, C. coniocraea and C. ochrochlora are included in the complex. In addition, C. maxima, C. phyllophora and C. subchordalis are supported as monophyletic species outside the C. gracilis complex. Cladonia maxima may form a separate species and variation among podetial morphology may be explained by convergent evolution.

Lichens Collected on an Alaska Highway Expedition in Alaska and Canada

A total of 622 species and 10 subspecies and varieties of lichens, plus six species of lichenicolous fungi, are reported from Alaska, the Yukon, British Columbia, and Alberta, mainly along the Alaska Highway and some of its side roads in lichenologically badly known regions. Anaptychia bryorum is new to the North American Arctic, Gongylia nadvornikii, Lichenodium sirosiphoideum and Stereocaulon capitellatum are new to North America, and Degelia plumbea is new to western North America. Verrucaria lobata Thomson sp. nov. is described from British Columbia. New to Alaska are: Acarospora superfusa, Amygdalaria subdissentiens, Aspicilia ka- relica, A. quartzitica, Bacidia herbarum, Brigantiaea fuscolutea, Bryonora curvescens, Bryoria friabilis, Calicium abietinum, Caloplaca borealis, C. ammiospila, C. sinapisperma, Chaenotheca xyloxena, Cladonia macroceras, C. ochrochlora, Collema crispum, C. curtisporum, Cyphelium tigillare, Graphis scripta, Heterodermia speciosa, Hypocenomyce scalaris, Hypogymnia occiden- talis, Lecanora fuscescens, Lecidea erythrophaea, L. leptoboloides, L. lulensis, L. stenotera, Le- procaulon subalbicans, Leptogium cyanescens, L. teretiuscula, Melanelia sorediata, M. tominii, Nephroma helveticum subsp. sipeanum, Parmelia hygrophila, Parmeliella triptophylla, Pertusaria borealis, Phylliscum tenue, Physcia subtilis, Psora lurida, Psorotichia schaereri, Pyrenopsis poly- cocca, Usnea filipendula, U. wasmuthii, Verrucaria funckii, Xanthoparmelia somloensis. New to Alberta are: Biatora carneoalbida, Bryoria capillaris, Candelariella lutella, C. xanthostigma, Collema furfuraceum, Lecania dubitans, Lecidea albohyalina, L. myriocarpella, Phaeophyscia cernohorskyi, P. hispidula, Ramalina obtusata, Rinodina oregona. New to British Columbia are: Acarospora smaragdula, A. veronensis, Actinogyra muehlenbergii, Adelolechia pilati, Arthror- haphis alpina, Aspicilia anseris, A. caesiopruinosa, A. disserpens, A. fimbriata, A. ryrkaipiae, A. subplicigera, Baeomyces roseus, Biatora carneoalbida, Bryocaulon divergens, Bryonora castanea, Caloplaca ammiospila, C. tetraspora, Catapyrenium cinereum, C. compactum, Cladonia thom- sonii, Collema glebulentum, Kiliasia athallina, Lecidea alpestris, Leciophysma finmarkicum, Le- praria lobificans, Micarea crassipes, Pertusaria alaskensis, P. coriacea, P. gyalectina, Phaeophyscia hirtella, Polyblastia obsoleta, P. terrestris, P. theleodes, Rhizocarpon eupetraeum, R. eupetraeoides, P. umbilicatum fo. pseudospeireum, Rinodina occidentalis, Staurothele drummondii, Thelidium absconditum, Toninia tristis, Usnea angulata, U. capitata, Verrucaria devergens, V. rupestris. New to Yukon are: Actinogyra muehlenbergii, Amygdalaria panaeola, Aspicilia aliena, A. cingulata, A. disserpens, A. heteroplaca, A. ryrkaipiae, A. subplicigera, Bryonora castanea, Bryoria trichodes subsp. americana, Buellia elegans, Calicium viride, Caloplaca crenularia, C. discolor, C. invadens, C. nivalis, C. pyracea, Candelariella athallina, C. dispersa, C. lutella, Cetraria califomica, Cladonia alaskana, C. bacilliformis, C. borealis, C. luteoalba, C. sulphurina, Collema crispum, Dermato- carpon intestiniforme, D. miniatum, Ephebe hispidula, Lecanora atrosulphurea, L. boligera, L. gangaleoides, L. intricata, L. straminea, L. strobilina, Lecidea alpestris, L. lapicida fo. ochracea, L. subcandida, Lecidella carpathica, L. elaeochroma, L. wulfenii, Lepraria caesioalba, Leproloma cacuminum, Leptogium tenuissimum, Melanelia infumata, Micarea peliocarpa, Parmeliella arc- tophila, Peltigera kristinssonii, Polyblastia sendtneri, Psilolechia lucida, Psorotichia schaereri, Rhizocarpon alpicola, R. cinereonigrum, Rinodina bischoffii, Squamarina lentigera, Staurothele elenkinii, Stereocaulon saxatile.

The checklist of parmelioid and similar lichens in Europe and some adjacent territories: additions and corrections

Abstract: Additions and corrections to the checklist published in 2008 are presented. Ten additionalspeciesarerecognized: Allantoparmeliaalmquistii,Asahineachrysantha,Hypogymniabryophila,Hypotra-chyna afrorevoluta, Melanelixia glabratula , Parmelia encryptata , Parmelina atricha , P. cryptotiliacea , Parmotrema cinereopruinata ,and Punctelia rudecta . Myelochroa subaurulenta is deleted, and Xanthopar-melia perezdepazii is also noted as newly described from the Canary Islands. In addition, Cavernulariahultenii is now placed in Hypogymnia ,the Canoparmelia crozalsiana group is included within Parmo-trema ,and Parmelinopsis is treated as a synonym of Hypotrachyna . The typification of the name Lichenquercinus is also corrected with the designation of the original figure as lectotype, and the newcombination Hypotrachyna cryptochlora comb. nov. (syn. Parmelia cryptochlora )ismade. Key words: Ascomycota , biogeography, Hypotrachyna cryptochlora , Lecanoromycetes, Parmeliaceae Introduction Hawksworth

Chemotyping glucans from lichens of the genus Cladonia

Abstract α- d -Glucans were isolated from the Cladonia spp., C. clathrata, C. connexa, C. crispatula, C. furcata, C. ibitipocae, C. imperialis, C. penicillata, and C. signata, to evaluate their possible significance in chemotyping. Each was isolated in water-insoluble form via successive alkaline extraction and freeze-thawing. They were then individually investigated using 13C and 1H-NMR spectroscopy (including COSY, TOCSY and HMQC techniques), methylation analysis, and Smith degradation, and their specific rotations and monosaccharide compositions determined. Each α- d -glucan contained alternating (1→3) and (1→4)-linkages (1:1 ratio), as determined by the relative areas of their respective H-1 signals at δ 5.14 and 5.23. Also, the respective relative areas of C-1 signals at δ 99.2 and 100.1, and those of C-3 of 3-O-(δ 82.6) and C-4 of 4-O-substituted (δ 78.9) units, were similar for each glucan. The presence of α- d -glucans of the nigeran type appears to be typical of Cladonia spp.

Nomenclatural Notes on Cladonia Species

The nomenclature and in part also the taxonomy of the following European lichens is discussed: Cladonia cervicornis (Ach.) Flotow subsp. cervicornis and subsp. verticillata (Hoffm.) Ahti comb. nov., C. bacilliformis (Nyl.) Gluck, C. portentosa (Dufour) Coem., C. anomaea (Ach.) Ahti & P. James comb. nov. (syn. C. pityrea (Florke) Fr.), C. conista Robbins ex Allen, C. conoidea Ahti sp. nov. (syn. C. conistea sensu Asah.: contains atranorin and fumarprotocetraric acid, type from England), C. bacillaris Nyl., C. coniocraea auct., and C. ochrochlora Florke. The African lichen ‘ Cladonia gorgonina (Bory) Eschw.’ is typified and its nomenclatural status reconsidered. Under the valid name Cladonia gorgonea Eschw. it is reduced to the synonymy of Cladia aggregata (Swartz) Nyl. The correct name for Cladonia gorgonina auct. is C. glaucopallida Vainio. A list of the new lichen taxa proposed by J. B. Bory, which were invalidly published by Florke in 1809, is also presented.

Phylogeny and taxonomy of the ‘manna lichens’

We present ataxonomic revision of the ‘manna lichens’ based on morphological, chemical, ecological and molecular data. A large number of herbarium specimens and fresh collections were examined. Phylogenetic analyses were performed using nuclear ribosomal (nrITS, nrLSU) and mitochondrial small subunit (mtSSU) sequences. Some notable phenotypic characters were plotted on the phylogenetic tree, and the analysis reveals that some of these characters are useful for genus and species level distinction of certain ‘manna lichens.’ Phylogeny of the Megasporaceae was revised using a combined data set of nrLSU and mtSSU and performing parsimony and Bayesian analyses. Five genera (Aspicilia, Circinaria, Lobothallia, Megaspora and Sagedia) are recognized. Further, the relationships of five presumably closely related genera of ‘manna lichens’, namely Agrestia (vagrant), Aspicilia (crustose) Circinaria (crustose), Chlorangium (vagrant) and Sphaerothallia (vagrant) with different growth forms were analysed. The analyses revealed that ‘manna lichens’ do not form a monophyletic group but occur in different clades within the genus Circinaria. The genera Agrestia, Chlorangium and Sphaerothallia are assigned as new synonyms under the genus Circinaria and no vagrant or erratic species remain in the genus Aspicilia. The analyses also show that five new erratic, vagrant and crustose species can be recognized. In this study two ‘manna lichens’, viz. Circinaria rostamii sp. nov. (Azerbaijan, Iran and Turkey), and Circinaria gyrosa sp. nov. (Armenia, Azerbaijan, Iran, Turkey, Turkmenistan and Spain) are described as new to science. Three potentially new species with crustose and erratic forms need additional study. Aspicilia fruticolosofoliacea is reduced to synonymy under C. alpicola, and a lectotype is designated for C. aspera. Thirteen new combinations in Circinaria are presented. The phenomenon of vagrancy is briefly discussed, and the biogeography of the ‘manna lichens’ is outlined. Illustrations, distribution maps, and an identification key to the species are provided.