Ulf Arup

A new taxonomy of the Caloplaca citrina group in the Nordic countries, except Iceland

The Caloplaca citrina group in the Nordic countries, except Iceland, is analysed with regard to morphology, anatomy, ecology, and chemistry. Their genetic relationship is also analysed using the ITS ribosomal DNA gene. The results show that there are at least five species within what has been called C. citrina, of which four are closely related to one another and to several non-sorediate species. Three of these, C. citrina, C. flavocitrina and C. arcis have previously been recognized at specific or varietal level some time during the last two centuries, whereas the fourth species, C. dichroa, is described as new. The fifth species, C. phlogina, is not related to the other species but shows a close relationship to the Xanthoria candelaria group. The species are described and illustrated in colour. Distribution maps are given for the study area and a key to sorediate, yellow to orange species occurring in the Nordic countries, except Iceland, is also provided.

The importance of host tree age, size and growth rate as determinants of epiphytic lichen diversity in boreal spruce forests

The amount of large and old trees has decreased in the boreal forests during the last centuries of forestry. Such trees are important habitats for epiphytic lichens and there is a growing concern for lichen species that are associated with large and old trees. However, only little is known about the relative importance of tree size versus age as determinants of lichen diversity. Here we have determined the size, age and growth rate of 157 Norway spruce trees and recorded the occurrence of epiphytic lichen species on their branches and lower stems. The study includes crustose lichens and was done in two old-growth forests in SE Norway. Tree age and tree size were the parameters that explained the largest part of epiphytic lichen diversity. Only the growth rate of the most recent time period, i.e. 1984–2004, showed a statistically significant relationship to diversity. There was no indication of a stabilising species number with increasing tree age. Slow-growing and old trees were, however, mainly of importance to the lichen species growing on stems, and this set of species were in general adversely affected by a large amount of branches. The opposite was the case for the species that were confined to branches as their diversity increased when the amount of branches increased. Our study adds empirical data to support the importance of large and old trees as bearers of biodiversity in boreal forests. Site preservation and patch retention of groups of old and large trees is recommended as measures to maintain epiphytic lichen diversity.

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.

MOLECULAR SYSTEMATICS OF LECANORA SUBGENUS PLACODIUM

A first hypothesis for the phylogeny of Lecanora subgen. Placodium is presented by using molecular data. Previous evolutionary ideas and classification concepts for this group with non-molecular data are re-investigated using DNA sequence data from the nuclear ITS and 5.8S regions. Using Protoparmelia as an outgroup, the Lecanora subfusca group together with the L. rupicola group appear as a sister group to assemblages with lobate species. Subgen. Placodium, as currently accepted, is not monophyletic. Molecular data suggest that the Lecanora dispersa group and the L. polytropa group are widened by lobate species and there is evidence from the ITS data that the monotypic genus Arctopeltis Poelt is closely related to the L. dispersa group. A congruence between molecular data and secondary chemistry supports the broader concept of the L. dispersa group suggested by the molecular data.

A phylogenetic study of the Lecanora rupicola group (Lecanoraceae, Ascomycota)

A molecular phylogeny of the Lecanora rupicola group is presented, based on ITS sequence analyses. The study includes saxicolous and corticolous members of the Lecanora rupicola group as well as other Lecanora species with pruinose apothecia. A phylogenetic hypothesis for species in Lecanora s. lat. and various other genera in Lecanoraceae, based on an alignment-free distance estimation technique, shows that the Lecanora rupicola group forms a monophyletic clade within Lecanoraceae. Affinities to the core group of Lecanora are not well supported, likewise the monophyly of Lecanora s. str. with other species groups in Lecanora, such as the lobate taxa (and Rhizoplaca) is not supported. A more detailed analysis involving Lecanora species with pruinose apothecial discs was carried out with model-based Bayesian Markov chain Monte Carlo (B/MCMC) tree sampling. The results suggest the monophyly of the Lecanora species that are characterized by the presence of chromones. Corticolous as well as saxicolous species are included. Lepraria flavescens is closely related to the Lecanora swartzii subgroup, and the new name Lecanora rouxii nom. nov. is introduced for that species. Other Lecanora species with pruinose discs are not closely related to the Lecanora rupicola group.

Taxonomy and phylogeny of the Caloplaca cerina group in Europe

Using ITS nrDNA sequence data, the Caloplaca cerina group (Teloschistaceae) is defined here as a monophyletic, but internally richly branched lineage. The group is also characterized by a combination of morphological and anatomical characters. Its internal lineages are supported by phenotypic characters in addition to ecology and distribution. Within the large C. cerina group, we have found at least 20 phylospecies in the temperate zone of the Northern Hemisphere. Two species complexes do not produce any vegetative diaspores: the polyphyletic, corticolous Caloplaca cerina s. lat. (six separated cryptic or semi-cryptic species) and the monophyletic C. stillicidiorum s. lat. that grows mainly on plant debris, small shrubs and bryophytes and consists of at least four internal lineages. All lineages producing vegetative diaspores (soredia, blastidia, isidia or lobules) are phenotypically characteristic and represent fairly easily distinguishable species: C. chlorina, C. isidiigera, C. monacensis, C. subalpina, C. thracopontica, C. turkuensis and C. ulmorum. Only the North American sorediate C. pinicola possibly represents an aggregate of species. Caloplaca sterilis is described as a new species. A key to the phenotypically distinguishable species is provided. Lectotypes are designated for C. albolutea, Caloplaca cerina f. coronulata and for C. monacensis. The Australian C. hanneshertelii belongs to this group, but this and other possible species from the Southern Hemisphere are not treated here in detail. Some species traditionally placed in the C. cerina group due to their similar morphology are excluded here on the basis of our phenotype examinations and molecular data. Caloplaca albolutea, C. mydalaea and C. virescens are uncertain taxa and their identities still remain unclear. (Less)

An exceptional group-I intron-like insertion in the SSU rDNA of lichen mycobionts

Abstract An exceptional group-I intron-like insertion at position 940 of the nuclear small subunit rDNA is found in lichen mycobionts of the families Parmeliaceae and Lecanoraceae. This shared insertion site is exceptional as it follows a G. Although several features of the self-splicing Tetrahymena intron are missing, the conserved structure of the presumed core region indicates that the new intron-like insertion, which is missing in mature transcripts, is not part of a silenced ribosomal repeat. It is unlikely that the new insertion is horizontally transferred from the adjacent position 943. A phylogenetic analysis indicates congruence with lichen phylogeny and suggests that the insertion has been vertically inherited.

The lichen genus Caloplaca (Ascomycota, Lecanoromycetes) on Svalbard. Notes and additions.

23 species of the lichen genus Caloplaca from Svalbard are described and/or discussed. The descriptions are natural language descriptions based on characters for each species coded into LIAS (Global Information System for Lichenized and Non-Lichenized Ascomycetes). A total of 37 Caloplaca species are listed for Svalbard, of which two species: C. elvebakkiana and C. scabrosa are described as new to science. C. alaskensis is recorded for the first time from Svalbard. A key to the Caloplaca species known from Svalbard is provided.

The Genus Caloplaca on Seashore Rocks in Eastern North America

The species ofthe genus Caloplaca occurring on seashore rocks in eastern North America have been studied with regard to taxonomy, geographical distribution, and ecology. Five species were found to inhabit the coastal rocks. The species also occur in Europe, and three of them in Asia. Three of the species, C. microthallina (Wedd.) Zahlbr., C. scopularis (Nyl.) Lettau, and C. verru- culifera (Vainio) Zahlbr. occur exclusively on seashore rocks, while C. lithophila H. Magn. and C. fraudans (Th. Fr.) H. Olivier occur also in inland localities. One-way analysis of variance of both morphological and anatomical characters was carried out on the American and the European populations of the species, except C. fraudans. No significant differences were found in C. lithophila, but in the other three species significant differences were recorded in two or three characters. However, the differences were mostly very small, the overlaps large, and no discontinuities could be detected. Therefore no infraspecific taxa were recognized. The lichen flora occurring on seashore rocks in eastern North America has previously been studied on several occasions. In the late 19th and the early 20th century contributions to the knowledge of lit- toral lichens were made mainly with scattered col- lections by botanists such as E. Tuckerman, H. Wil- ley, W. G. Farlow, and G. K. Merrill. Degelius (1940) was one of the first lichenologists to investigate the lichens on seashore rocks in North America more thoroughly. He studied the flora of Maine and com- pared it with the European flora, and found great similarities. The work of Degelius inspired I. M. Lamb (1954) to carry out similar studies in Nova Scotia, Canada. The most extensive collections of littoral lichens in eastern North America were made by R. M. Tay- lor. His work mainly concerned the vertical distri- bution patterns, interspecific associations, and the general geographical distribution of the species col- lected, but also comprised taxonomical studies of the genus Verrucaria and a key to the 21 species included in the study (Taylor 1974). One of the latest studies in eastern North America, including littoral lichens, is that of the flora of Fundy National Park, New Brunswick, Canada, by Gowan and Bro- do (1988). They pointed out that the maritime prov- inces are still not very well known and that the need for further studies is high. This study was carried out to investigate the spe- cies of Caloplaca occurring on seashore rocks in eastern North America, and to compare the Euro- pean and the American populations of the species. No extensive studies have previously been done on this group in this region. Three of the species, oc- curring also in northern Europe, have been treated by Nordin (1972), but only for the European range of the species.

Towards a revised generic classification of lecanoroid lichens (Lecanoraceae, Ascomycota) based on molecular, morphological and chemical evidence

The phylogenetic relationship of lecanoroid lichens is studied using two data sets: 1) a 2-locus data set including 251 OTUs representing 150 species, and 2) a 6-locus data set with 82 OTUs representing 53 species. The genus Lecanora as currently circumscribed is shown to be highly polyphyletic and several genera, including Adelolecia, Arctopeltis, Bryonora, Carbonea, Frutidella, Lecidella, Miriquidica, Palicella, Protoparmeliopsis, Pyrrhospora, and Rhizoplaca are nested within Lecanora sensu lato. A core group of Lecanora is supported as monophyletic and includes species of the L. carpinea, L. rupicola, and L. subcarnea groups, and a core group of the L. subfusca group. Three monophyletic clades that are well supported in our analyses and well characterized by phenotypical characters are accepted here: 1) Myriolecis to accommodate the Lecanora dispersa group and Arctopeltis; 2) Protoparmeliopsis for the L. muralis group; and 3) Rhizoplaca is emended to include three placodioid taxa previously classified in Lecanora (L. novomexicana. L. opiniconensis, L. phaedrophthalma), whereas R. aspidophora and R. peltata are excluded from Rhizoplaca. The latter is transferred into Protoparmeliopsis. Lecidella is strongly supported as a monophyletic group. Our studies indicate the presence of additional clades of species currently placed in Lecanora sensu lato that warrant taxonomic recognition but additional data will be necessary before the circumscription of these entities is fully understood. 37 new combinations are proposed into the genera Myriolecis (30), Protoparmeliopsis (2), and Rhizoplaca (5).