Filip Högnabba

Implementing a cumulative supermatrix approach for a comprehensive phylogenetic study of the Teloschistales (Pezizomycotina, Ascomycota)

The resolution of the phylogenetic relationships within the order Teloschistales (Ascomycota, lichen-forming-fungi), with nearly 2000 known species and outstanding phenotypic diversity, has been hindered by the limitation in the resolving power that single-locus or two-locus phylogenetic studies have provided to date. In this context, an extensive taxon sampling within the Teloschistales with more loci (especially nuclear protein-coding genes) was needed to confront the current taxonomic delimitations and to understand evolutionary trends within this order. Comprehensive maximum likelihood and bayesian analyses were performed based on seven loci using a cumulative supermatrix approach, including protein-coding genes RPB1 and RPB2 in addition to nuclear and mitochondrial ribosomal RNA-coding genes. We included 167 taxa representing 12 of the 15 genera recognized within the currently accepted Teloschistineae, 22 of the 43 genera within the Physciineae, 49 genera of the closely related orders Lecanorales, Lecideales, and Peltigerales, and the dubiously placed family Brigantiaeaceae and genus Sipmaniella. Although the progressive addition of taxa (cumulative supermatrix approach) with increasing amounts of missing data did not dramatically affect the loss of support and resolution, the monophyly of the Teloschistales in the current sense was inconsistent, depending on the loci-taxa combination analyzed. Therefore, we propose a new, but provisional, classification for the re-circumscribed orders Caliciales and Teloschistales (previously referred to as Physciineae and Teloschistineae, respectively). We report here that the family Brigantiaeaceae, previously regarded as incertae sedis within the subclass Lecanoromycetidae, and Sipmaniella, are members of the Teloschistales in a strict sense. Within this order, one lineage led to the diversification of the mostly epiphytic crustose Brigantiaeaceae and Letrouitiaceae, with a circumpacific center of diversity and found mostly in the tropics. The other main lineage led to another epiphytic crustose family, mostly tropical, and with an Australasian center of diversity--the Megalosporaceae--which is sister to the mainly rock-inhabiting, cosmopolitan, and species rich Teloschistaceae, with a diversity of growth habits ranging from crustose to fruticose. Our results confirm the use of a cumulative supermatrix approach as a viable method to generate comprehensive phylogenies summarizing relationships of taxa with multi-locus to single locus data.

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.

High selectivity in symbiotic associations of lichenized ascomycetes and cyanobacteria

The selectivity of mycobionts and cyanobionts in lichen symbioses were examined. We analyzed symbiotic cyanobionts, collected from different sample sites, and compared them to free‐living cyanobacteria Nostoc. Cyanobionts were obtained from lichens assigned to the genera Pseudocyphellaria and Sticta, in particular. Multiple gene loci were screened and direct optimization was used in the phylogenetic analyses. We show that many lichen fungi are strongly selective towards their cyanobionts. Lichenized ascomycetes seem to be able to identify and choose a specific strain, species or a species group of Nostoc with which to associate. The present analyses also suggest that some of the Nostoc taxa may be specialized in symbiotic life with only lichenized ascomycetes. Despite the selectivity observed in fungi, there appears to be no coevolution between the partners. We have also discussed the problems of using the tRNALeu intron as a marker in phylogenetic analyses.

Molecular phylogeny of the Sphaerophorus globosus species complex

The Sphaerophorus globosus complex (Lecanorales, lichenized Ascomycota) shows a large morphological variation, and three relatively distinct morphotypes can be distinguished in parts of the distribution area. Here, we utilize a multigene‐based maximum‐parsimony approach (nITS+ LSU rDNA, mtSSU rDNA, β‐tubulin, and actin) to investigate whether these morphotypes constitute distinct species. The results show that there are at least two well‐supported monophyletic groups that we interpret as phylogenetic species within the S. globosus complex. These species do not completely correspond to the predefined morphotypes. One group, an apparently undescribed species, contains noncoralloid specimens from the North American Pacific Northwest. The other group, S. globosus, consists of two well‐supported monophyletic groups: one contains coralloid epiphytic specimens from the North American Pacific Northwest that are morphologically indistinguishable from epiphytic specimens from Europe and are presently interpreted as belonging to the same species and the other is morphologically variable and contains terrestrial specimens from Europe, North America, and southernmost South America and coralloid epiphytic and epilithic specimens from Europe. The results suggest that the population in southernmost South America originated by long‐distance dispersal from arctic populations in the Northern Hemisphere.

Phylogenetic relationships of Stereocaulaceae based on simultaneous analysis of beta-tubulin, GAPDH and SSU rDNA sequences

The phylogenetic relationships of the lichenised ascomycete family Stereocaulaceae were investigated with partial beta-tubulin, partial glyceraldehyde 3-phosphate dehydrogenase (GAPDH) and SSU rDNA sequence data. Due to the marked length variation in the SSU rDNA sequences, the analyses were performed with direct optimization. The combined analysis of the beta-tubulin, GAPDH and SSU rDNA sequences as well as the separate analyses of beta-tubulin and GAPDH data recognized the monophyletic group comprising Lepraria, Muhria and Stereocaulon. In the combined analysis Stereocaulon tornense appears as a basal taxon in the Stereocaulon clade, suggesting that the crustose growth form and the absence of cephalodia are primitive features within the genus. Muhria is nested in Stereocaulon and groups with the crustose S. cumulatum and S. leucophaeopsis. The genus Lepraria is monophyletic if Leproloma is included, but species that were represented by several specimens in our study, i.e., Lepraria jackii and Leproloma membranaceun, do not appear as monophyletic entities. The genus Pilophorus is included in Cladoniaceae. Combined analysis of the three data-sets clearly led to an increase in support values compared to the separate analyses, showing high support for the monophyly of families Cladoniaceae and Stereocaulaceae and for the sistergroup relation for the two families. The relationships between higher groupings did not reach high support values probably due to the incongruencies among the three datasets. (Less)

Expansion of the Stictidaceae by the addition of the saxicolous lichen-forming genus Ingvariella

The monotypic, lichen-forming genus Ingvariella originally was segregated from Diploschistes and placed within the Thelotremataceae (Ostropales) based on aspects of exciple morphology. However, the I+ hymenium and amyloid ascus wall suggest affinities to families other than the Thelotremataceae. To assess the identity of Ingvariella and to investigate its placement within the Ostropales, we inferred phylogenetic relationships of I. bispora by comparison of mtSSU rDNA and nuLSU rDNA sequences for 59 species encompassing a broad array of ostropalean fungi by means of Bayesian, maximum likelihood and weighted maximum parsimony methods. Here we report that Ingvariella is a member of the Stictidaceae, sister to the mainly saprotrophic genus Cryptodiscus. The inclusion of the first saxicolous lichen-forming fungus within this family expands the broad ecological diversity of the Stictidaceae, where saprotrophic fungi, corticicolous lichen-forming fungi and lichenized and non-lichenized conspecific taxa have been described previously. We also present new insights into the relationships among other families within the Ostropales.

High fungal selectivity for algal symbionts in the genus Bryoria

In this study we examined photobiont identity, diversity and selectivity in the genus Bryoria . We focused on B. fremontii and section Implexae in order to determine whether secondary chemistry is correlated with photobiont identity. DNA from two loci for photobionts and three loci for mycobionts was sequenced for both parsimony and Bayesian phylogenetic analyses. A comparison of photobiont and mycobiont phylogenies reveals that most Bryoria species associate exclusively with lineages of the Trebouxia simplex group; only B. smithii was associated with a different photobiont. We conclude that most Bryoria species included in our study are highly selective in their choice of algal partners and that the presence/concentration of different secondary compounds does not correlate with photobiont identity either in section Implexae or in B. fremontii .

Aspicilia rogeri sp. nov. (Megasporaceae) and other allied vagrant species in North America

Abstract A short revision of the vagrant Aspicilia species of North America is presented based on morphological, molecular and ecological data. Vagrant Aspicilia are common lichens throughout the steppes of the western United States and in southwestern parts of Canada. Species delimitation of these lichens is difficult because of the paucity of morphological characters and large degree of variation. Inferences from nuITSrDNA sequences reveals that the North American specimens of A. fruticulosa are not most closely related to their Eurasian populations but instead share a unique ancestor with A. hispida. The specimens of A. fruticulosa from the New World are hereby recognized as a distinct species, A. rogeri. Its differentiation from the similar A. fruticulosa and A. hispida is discussed. The exclusion of A. fruticulosa from the N. American checklist is proposed temporarily.

Phylogenetic position of the crustose Stereocaulon species

Phylogenetic relationships of Stereocaulon with emphasis on the crustose taxa were studied based on nuclear ribosomal ITS1-5.8S-ITS2 and partial beta-tubulin sequences. The placement of four of the six crustose species currently included in the genus has previously been confirmed based on molecular data. It has, however, remained unresolved whether the crustose growth form is a plesiomorphic or apomorphic feature within Stereocaulon, due to contradictory placements of the crustose species in earlier studies. The aim of this study was to clarify the position of the crustose species by including additional data, especially of S. nivale and S. plicatile, which have not been included in previous analyses. The inclusion of S. plicatile in the genus is of particular interest as it is the only species in the genus with submurifrom to muriform ascospores. Altogether 37 specimens representing 31 species of the ingroup, including all the crustose Stereocaulon species, were incorpo- rated in the analyses. Conventional, as well as direct optimization parsimony, maximum likelihood and Bayesian analyses were performed. The results show that the crustose species do not form a monophyletic entity and that the crustose growth form is a plesiomorphic feature within Stereocaulon. The crustose S. nivale and S. plicatile are nested within the genus and their inclusion in Stereocaulon is thereby confirmed. The nested position of S. plicatile indicates that the submuriform to muriform spore type has been gained independently within the genus. Here, S. plicatile is also reported for the first time from Scandinavia.

Lichens, Lichenicolous and Allied Fungi Found in Asveja Regional Park (Lithuania)

Abstract Motiejūnaitė J., Berglund T., Czarnota P., Himelbrant D., Högnabba F., Konoreva L. A., Korchikov E. S., Kubiak D., Kukwa M., Kuznetsova E., Leppik E., Lõhmus P., Prigodina Lukošienė I., Pykälä J., Stončius D., Stepanchikova I., Suija A., Thell A., Tsurykau A., Westberg M., 2012: Lichens, lichenicolous and allied fungi found in Asveja Regional Park (Lithuania) [Kerpės, lichenofiliniai ir kerpėms artimi saprotrofiniai grybai Asvejos regioniniame parke]. - Bot. Lith., 18(2): 85-100. The paper reports the results of lichenological investigations in Asveja Regional Park (eastern Lithuania). A large part of the study was performed during the joint 18th Symposium of the Baltic Mycologists and Lichenologists (BMLS) and Nordic Lichen Society (Nordisk Lichenologisk Förening, NLF) meeting on 19-23 September 2011. A list of 259 species is presented. Of these, 30 species are new to Lithuania. Arthonia helvola, Bacidina sulphurella, Candelariella lutella, Catillaria croatica, Cladonia conista, Gyalecta derivata, Lecanoraquercicola, Leptosphaeria ramalinae, Strigula jamesii, Trichonectria rubefaciens, Verrucaria banatica, V. boblensis, V. christiansenii, V. illinoisensis, V. inornata, V. nigrofusca, V. trabicola, Zwackhiomyces diederichii were recorded for the first time in the Baltic countries. New lichens to Lithuania are as follows: Bacidiaincompta, Caloplaca crenulatella, C. pyracea, Catinaria atropurpurea, Lecanora populicola, L. semipallida, Mycobilimbia epixanthoides, Ramalina dilacerata, Verrucaria inaspecta, and new lichenicolous fungi are: Cladosporium licheniphilum, Stigmidium microspilum, Xenonectriella leptalea. Eighteen species included in the Lithuanian Red Data Book were recorded, which is the highest number known for any studied area in Lithuania.