Ingvar Kärnefelt

Monophyletic groups within the Parmeliaceae identified by ITS rDNA, beta-tubulin and GAPDH sequences

Phylogenetic relationships within the Parmeliaceae are analysed cladistically on the basis of DNA characters from partial β-tubulin, partial glyceraldehyde-3-phosphate dehydrogenase (GAPDH) and ITS sequences. 100 taxa representing 73 of the 88 genera currently recognised are included in the analyses. Eight monophyletic groups including two or more genera were identified in the tree calculated from the combined data matrix. Three of the groups cover almost half of the species of the family. The largest and strongest supported group includes seven genera with their distribution centres in the Southern Hemisphere: Almbornia, Chondropsis, Karoowia, Namakwa, Neofuscelia, Xanthomaculina and Xanthoparmelia. The second group is a clade of four essentially tropical genera: Concamerella, Flavoparmelia, Parmotrema and Rimelia. The third large group with strong support is the core of cetrarioid lichens, distributed primarily in cold areas of the Northern Hemisphere. The genus Parmelia sensu Hale is not closely related with most of its segregates. One new combination, Cetrariella commixta, is proposed. Coelopogon abraxas is reported from South America for the first time.

Phylogeny of cetrarioid lichens (Parmeliaceae) inferred from ITS and beta-tubulin sequences, morphology, anatomy and secondary chemistry.

Phylogenetic relationships within the family Parmeliaceae (lichenized ascomycetes) with emphasis on the heterogeneous group of cetrarioid lichens are reconstructed. The results are based on cladistic analyses of DNA-sequences, morphological and chemical data. Almost all currently recognized cetrarioid genera were included in the analyses together with parmelioid and alectorioid members of the presumably monophyletic family Parmeliaceae. We tried to sample taxonomic diversity of the family as widely as possible. The ITS1-5.8S-ITS2 region of the rDNA and a partial β-tubulin gene from 126 samples representing 82 species were analysed. Cetrarioid lichens were identified as a monophyletic group, supported by both ITS and β-tubulin characters. This group was reanalysed using 47 morphological, anatomical and secondary chemistry characters combined with the DNA data matrix. ITS and β-tubulin sequences provide congruent information, and a clear correlation between DNA-data and conidial shape is observed. The current taxonomy of the cetrarioid lichens is discussed and compared with the phylogenetic trees obtained here. A comprehensive study of the phylogeography of some bipolar or subcosmopolitic species with representatives from both hemispheres was performed. Cetraria aculeata is the only taxon where correlation between DNA-data and geographic origin is observed.

Evolution and phylogeny of cetrarioid lichens

TheAcharian genusCetraria has not been scrutinised in the same way as the genusParmelia in terms of circumscription of segregate genera. A few generic names have been introduced, but mainly in checklists without any real indication of what these names stand for. After a detailed investigation of characters in the apothecia, conidiomata, anatomical structures, as well as morphology and secondary chemistry, it is clear that several, more or less distinctive, phylogenetic lines exist among taxa which earlier were accommodated inCetraria. Some of these distinctive phylogenetic lines are distinguished by ascus characters: (1) clavate asci with a small axial body, an apical ring structure in the tholus and ellipsoid ascospores; (2) uniseriate asci with a large axial body and globose ascospores; and (3) taxa with broadly clavate asci with a large axial body and ellipsoid ascospores. Sixty-three character states were studied and results from the character analyses were evaluated by cladistic analysis against 43 examined terminal taxa. Seven further taxa were included in one analysis. Results from the cladistic analyses give rather strong support for a new systematic treatment of at least certain groups in the cetrarioid lichens. A formal systematic arrangement is not made here but will be presented elsewhere. The characters investigated are illustrated in 78 half tone pictures and one line drawing. Five cladograms are presented.

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.

Provitamins and vitamins D2 and D3 in Cladina spp. over a latitudinal gradient: possible correlation with UV levels

Provitamin D2, vitamin D2 and vitamin D3 were identified in the thallus of a lichen species, Cladina arbuscula (Wallr.) Hale and W.L. Culb. The identification of vitamin D3 was supported by: (1) co-chromatography in both reverse and straight phase HPLC (high performance liquid chromatography), (2) ultraviolet absorption spectrum, and (3) molecular ion peaks demonstrated by ESI (electrospray ionisation) mass spectrometry. The contents of vitamin D3 range from 0.67 to 2.04 μg g⁻¹ dry matter in the thalli of C. arbuscula specimens grown under different natural conditions, while provitamin D3 could not be detected. The ranges for provitamin D2 and vitamin D2 were 89-146 and 0.22-0.55 μg g⁻¹ dry matter, respectively, while the contents of provitamin D3 were below the detection limit (0.01 microg g(-1) dry matter). When C. arbuscula thalli collected at different latitudes from northern Finland to Greece were compared, a positive correlation of vitamin D2 and D3 contents with modelled UV-B radiation at the collection sites was found. A single sample of C. rangiferina from northern Finland gave much higher values for the vitamins. A possible reason could be the lower content of UV-B absorbing pigment in the latter species.

The evaluation of characters in lichenized families, exemplified with the alectorioid and some parmelioid genera

Important characters in the classification of the lichenized familyAlectoriaceae (Lecanorales) are discussed and evaluated. A few associated genera presently accommodated in the familyParmeliaceae (Lecanorales) are also discussed. Characters in the asci and hamathecium are of major importance, but structural characters such as general thallus organization and anatomy of cortical layers must also be considered. Secondary chemical products, however, were not found to be decisive for the circumscription of this family. Only three genera,Alectoria, Oropogon, andSulcaria, can be included, based mainly on the structure of the large, strongly amyloid asci; the rather large and darkly pigmented, thick-walled spores, usually few per ascus; and the structure of the anastomosing paraphysoids. Arguments are also given as to why the other alectorioid genera, i.e.Bryocaulon, Bryoria andPseudephebe must be excluded from this family. Similarly developed ascus and hamathecial structures are also discussed in a few unrelated parmelioid genera:Cetrelia, Menegazzia, Parmelaria, andParmelia.

The lichen genera Arctocetraria, Cetraria, and Cetrariella (Parmeliaceae) and their presumed evolutionary affinities

The lichen genera Arctocetraria, Cetraria, and Cetrariella (Parmeliaceae) and their presumed evolutionary affinities

What is Caloplaca cinnabarina

This paper treats four saxicolous species of Caloplaca occurring mainly in subtropical or semiarid regions in North and Central America, C. baueri, C. cinnabarina, C. rosei, and C. rubelliana. These species are mainly characterized by 1) a continuous to cracked areolate thallus, 2) thallus without marginal lobes, 3) areole margins broadly attached to the substrate, 4) cortex distinctly paraplectenchymatous, 5) immersed apothecia, and 6) containing anthraquinones. The group is not regarded as natural and consequently not given taxonomic status. Descriptions and distribution maps are provided for the taxa. Color illustrations or references to earlier published color illustrations are provided for all species. Additional notes are presented for two similar non American species, C. haematodes and C. subunicolor. (Less)

Isidiate entities in the Teloschistales and their ecological and evolutionary significance.

Thirteen isidiate taxa in the Teloschistales are briefly discussed, i.e. Letrouitia corallina (Mull. Arg.) Hafellner, L. leprolyta (Nyl.) Hafellner, Caloplaca bassiae (Ach.) Zahlbr., C. granulosa (Mull. Arg.) Jatta, C. herbidella (Hue) H. Magn., C. isidiigera Vězda, C. isidiosa (Vainio) Zahlbr., C. littorea Tavares, C. namibensis Karnef., C. ochraceofulva (Mull. Arg.) Jatta, C. verruculifera (Vainio) Zahlbr., Xanthoria domogledensis Vezda and X. isidioidea (Beltr.) Szat. The presumed origins and affinities of these taxa, which are also often fertile, are discussed. The role of isidia in reproduction and dispersal, and the justification of specific status in some of these taxa is questioned.

A new circumscription of the lichen genus Nephromopsis (Parmeliaceae, lichenized Ascomycetes).

The phylogeny of the cetrarioid lichens with bifusiform spermatia and dorsiventral thalli which contain usnic acid is reanalysed using three parts of the genome, ITS rDNA, β-tubulin and GAPDH sequences. Molecular data from five cetrarioid species are presented for the first time, and 13 new sequences are combined with sequences from the gene bank to delimit the genus Nephromopsis. A monophyletic clade of Nephromopsis, Tuckneraria, ‘Cetraria’ leucostigma and ‘C.’ melaloma is identified and circumscribed as one genus, Nephromopsis, which now includes 19 species. Four new combinations are presented. A key to the species is provided.