Lichens and associated fungi from Glacier Bay National Park, Alaska.

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 lichenicolous 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 alaskana, 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.

Biatoraalnetorum (Ramalinaceae, Lecanorales), a new lichen species from western North America.

Biatoraalnetorum S. Ekman & Tønsberg, a lichenised ascomycete in the family Ramalinaceae (Lecanorales, Lecanoromycetes), is described as new to science. It is distinct from other species of Biatora in the combination of mainly three-septate ascospores, a crustose thallus forming distinctly delimited soralia that develop by disintegration of convex pustules and the production of atranorin in the thallus and apothecia. The species is known from the Pacific Northwest of North America, where it inhabits the smooth bark of Alnusalnobetulasubsp.sinuata and A.rubra. Biatoraalnetorum is also a new host for the lichenicolous ascomycete Sclerococcumtoensbergii Diederich.

Two Basidiomycete Fungi in the Cortex of Wolf Lichens.

Since the late 1800s, mycologists have been detecting fungi above and beyond the assumed single fungus in lichen thalli [1-6]. Over the last century, these fungi have been accorded roles ranging from commensalists to pathogens. Recently, Cyphobasidiales yeasts were shown to be ubiquitous in the cortex layer of many macrolichens [7], but for most species, little is known of their cellular distribution and constancy beyond visible fruiting structures. Here, we demonstrate the occurrence of an additional and distantly related basidiomycete, Tremella, in 95% of studied thalli in a global sample of one of the most intensively studied groups of lichens, the wolf lichens (genus Letharia). Tremella species are reported from a wide range of lichen genera [8], but until now, their biology was deduced from fruiting bodies (basidiomata) formed on lichen thalli. Based on this, they have been thought to be uncommon to rare, to occur exclusively in a hyphal form, and to be parasitic on the dominant fungal partner [9, 10]. We show that, in wolf lichens, Tremella occurs as yeast cells also in thalli that lack basidiomata and infer that this is its dominant stage in nature. We further show that the hyphal stage, when present in Letharia, is in close contact with algal cells, challenging the assumption that lichen-associated Tremella species are uniformly mycoparasites. Our results suggest that extent of occurrence and cellular interactions of known fungi within lichens have historically been underestimated and raise new questions about their function in specific lichen symbioses.

Evolution of growth habit, inflorescence architecture, flower size, and fruit type in Rubiaceae: its ecological and evolutionary implications.

During angiosperm evolution, innovations in vegetative and reproductive organs have resulted in tremendous morphological diversity, which has played a crucial role in the ecological success of flowering plants. Morindeae (Rubiaceae) display considerable diversity in growth form, inflorescence architecture, flower size, and fruit type. Lianescent habit, head inflorescence, small flower, and multiple fruit are the predominant states, but arborescent habit, non-headed inflorescence, large flower, and simple fruit states occur in various genera. This makes Morindeae an ideal model for exploring the evolutionary appearances and transitions between the states of these characters. We reconstructed ancestral states for these four traits using a bayesian approach and combined nuclear/chloroplast data for 61 Morindeae species. The aim was to test three hypotheses: 1) self-supporting habit is generally ancestral in clades comprising both lianescent and arborescent species; 2) changes from lianescent to arborescent habit are uncommon due to "a high degree of specialization and developmental burden"; 3) head inflorescences and multiple fruits in Morindeae evolved from non-headed inflorescences and simple fruits, respectively. Lianescent habit, head inflorescence, large flower, and multiple fruit are inferred for Morindeae, making arborescent habit, non-headed inflorescence, small flower, and simple fruit derived within the tribe. The rate of change from lianescent to arborescent habit is much higher than the reverse change. Therefore, evolutionary changes between lianescent and arborescent forms can be reversible, and their frequency and trends vary between groups. Moreover, these changes are partly attributed to a scarcity of host trees for climbing plants in more open habitats. Changes from large to small flowers might have been driven by shifts to pollinators with progressively shorter proboscis, which are associated with shifts in breeding systems towards dioecy. A single origin of dioecy from hermaphroditism is supported. Finally, we report evolutionary changes from headed to non-headed inflorescences and multiple to simple fruits.

Phylogeny and character evolution in the jelly fungi (Tremellomycetes, Basidiomycota, Fungi).

The Tremellomycetes (Agaricomycotina, Basidiomycota, Fungi) are a nutritionally heterogeneous group comprising saprotrophs, animal parasites, and fungicolous species (fungal-inhabiting, including lichen-inhabiting). The relationships of many species, particularly those with a lichenicolous habit, have never been investigated by molecular methods. We present a phylogeny of the Tremellomycetes based on three nuclear DNA ribosomal markers (nSSU, 5.8S and nLSU), representing all main taxonomic groups and life forms, including lichenicolous taxa. The Cystofilobasidiales, Filobasidiales, Holtermanniales, and Tremellales (including the Trichosporonales) are recovered as monophyletic, but this is not the case for the Tremellomycetes. We suggest, however, that the Cystofilobasidiales tentatively continue to be included in the Tremellomycetes. As currently circumscribed, the Filobasidiaceae, Sirobasidiaceae, Syzygosporaceae and Tremellaceae are non-monophyletic. Cuniculitremaceae, Sirobasidiaceae and Tetragoniomycetaceae are nested within Tremellaceae. The lichenicolous species currently included within the Tremellomycetes belong in this group, distributed across the Filobasidiales and Tremellales. Lichen-inhabiting taxa do not form a monophyletic group; they are distributed in several clades and sometimes intermixed with taxa of other nutritional habits. Character state reconstruction indicates that two morphological traits claimed to characterize groups in the Tremellomycetes (the basidium habit and basidium septation) are highly homoplastic. Comparative phylogenetic methods suggest that the transitions between single and catenulate basidia in the Tremellales are consistent with a punctuational model of evolution whereas basidium septation is likely to have evolved under a graduational model in the clade comprising the Holtermanniales, Filobasidiales, and Tremellales.

The devil in the details: interactions between the branch-length prior and likelihood model affect node support and branch lengths in the phylogeny of the Psoraceae.

In popular use of Bayesian phylogenetics, a default branch-length prior is almost universally applied without knowing how a different prior would have affected the outcome. We performed Bayesian and maximum likelihood (ML) inference of phylogeny based on empirical nucleotide sequence data from a family of lichenized ascomycetes, the Psoraceae, the morphological delimitation of which has been controversial. We specifically assessed the influence of the combination of Bayesian branch-length prior and likelihood model on the properties of the Markov chain Monte Carlo tree sample, including node support, branch lengths, and taxon stability. Data included two regions of the mitochondrial ribosomal RNA gene, the internal transcribed spacer region of the nuclear ribosomal RNA gene, and the protein-coding largest subunit of RNA polymerase II. Data partitioning was performed using Bayes' factors, whereas the best-fitting model of each partition was selected using the Bayesian information criterion (BIC). Given the data and model, short Bayesian branch-length priors generate higher numbers of strongly supported nodes as well as short and topologically similar trees sampled from parts of tree space that are largely unexplored by the ML bootstrap. Long branch-length priors generate fewer strongly supported nodes and longer and more dissimilar trees that are sampled mostly from inside the range of tree space sampled by the ML bootstrap. Priors near the ML distribution of branch lengths generate the best marginal likelihood and the highest frequency of "rogue" (unstable) taxa. The branch-length prior was shown to interact with the likelihood model. Trees inferred under complex partitioned models are more affected by the stretching effect of the branch-length prior. Fewer nodes are strongly supported under a complex model given the same branch-length prior. Irrespective of model, internal branches make up a larger proportion of total tree length under the shortest branch-length priors compared with longer priors. Relative effects on branch lengths caused by the branch-length prior can be problematic to downstream phylogenetic comparative methods making use of the branch lengths. Furthermore, given the same branch-length prior, trees are on average more dissimilar under a simple unpartitioned model compared with a more complex partitioned models. The distribution of ML branch lengths was shown to better fit a gamma or Pareto distribution than an exponential one. Model adequacy tests indicate that the best-fitting model selected by the BIC is insufficient for describing data patterns in 5 of 8 partitions. More general substitution models are required to explain the data in three of these partitions, one of which also requires nonstationarity. The two mitochondrial ribosomal RNA gene partitions need heterotachous models. We found no significant correlations between, on the one hand, the amount of ambiguous data or the smallest branch-length distance to another taxon and, on the other hand, the topological stability of individual taxa. Integrating over several exponentially distributed means under the best-fitting model, node support for the family Psoraceae, including Psora, Protoblastenia, and the Micarea sylvicola group, is approximately 0.96. Support for the genus Psora is distinctly lower, but we found no evidence to contradict the current classification.

Slippery when wet: phylogeny and character evolution in the gelatinous cyanobacterial lichens (Peltigerales, Ascomycetes).

Many lichen fungi form symbioses with filamentous Nostoc cyanobacteria, which cause the lichen to swell and become extremely gelatinous when moist. Within the Lecanoromycetes, such gelatinous lichens are today mainly classified in the Collemataceae (Peltigerales, Ascomycota). We performed Bayesian MCMC, maximum likelihood, and maximum parsimony analyses of three independent markers (mtSSU rDNA, nuLSU rDNA, and RPB1), to improve our understanding of the phylogeny and classification in the Peltigerales, as well as the evolution of morphological characters that have been used for classification purposes in this group. The Collemataceae and the non-gelatinous Pannariaceae are paraphyletic but can be re-circumscribed as monophyletic if Leciophysma, Physma, Ramalodium and Staurolemma are transferred to the Pannariaceae. The gelatinous taxa transferred to the Pannariaceae deviate from other Collemataceae in having simple ascospores, and several also have a ring-shaped exciple as in other Pannariaceae, rather than the disc-shaped exciple found in the typical Collemataceae. Both Collema and Leptogium are non-monophyletic. The re-circumscribed Collemataceae shares a distinct ascus type with the sister group Placynthiaceae and the Coccocarpiaceae, whereas Pannariaceae includes a variety of structures. All Pannariaceae have one-celled ascospores, whereas all Collemataceae have two- or multi-celled spores. Reconstructions of the number of character state transformations in exciple structure, thallus gelatinosity, and ascus apex structure indicate that the number of transformations is distinctly higher than the minimum possible. Most state transformations in the exciple took place from a ring-shaped to a disc-shaped exciple. Depending on the reconstruction method, most or all transformations in thallus structure took place from a non-gelatinous to a gelatinous thallus. Gains and losses of internal structures in the ascus apex account for all or a vast majority of the number of transformations in the ascus, whereas direct transformations between asci with internal structures appear to have been rare.

The limitations of ancestral state reconstruction and the evolution of the ascus in the Lecanorales (lichenized Ascomycota).

Ancestral state reconstructions of morphological or ecological traits on molecular phylogenies are becoming increasingly frequent. They rely on constancy of character state change rates over trees, a correlation between neutral genetic change and phenotypic change, as well as on adequate likelihood models and (for Bayesian methods) prior distributions. This investigation explored the outcomes of a variety of methods for reconstructing discrete ancestral state in the ascus apex of the Lecanorales, a group containing the majority of lichen-forming ascomycetes. Evolution of this character complex has been highly controversial in lichen systematics for more than two decades. The phylogeny was estimated using Bayesian Markov chain Monte Carlo inference on DNA sequence alignments of three genes (small subunit of the mitochondrial rDNA, large subunit of the nuclear rDNA, and largest subunit of RNA polymerase II). We designed a novel method for assessing the suitable number of discrete gamma categories, which relies on the effect on phylogeny estimates rather than on likelihoods. Ancestral state reconstructions were performed using maximum parsimony and maximum likelihood on a posterior tree sample as well as two fully Bayesian methods. Resulting reconstructions were often strikingly different depending on the method used; different methods often assign high confidence to different states at a given node. The two fully Bayesian methods disagree about the most probable reconstruction in about half of the nodes, even when similar likelihood models and similar priors are used. We suggest that similar studies should use several methods, awaiting an improved understanding of the statistical properties of the methods. A Lecanora-type ascus may have been ancestral in the Lecanorales. State transformations counts, obtained using stochastic mapping, indicate that the number of state changes is 12 to 24, which is considerably greater than the minimum three changes needed to explain the four observed ascus apex types. Apparently, the ascus in the Lecanorales is far more apt to change than has been recognized. Phylogeny corresponds well with morphology, although it partly contradicts currently used delimitations of the Crocyniaceae, Haematommataceae, Lecanoraceae, Megalariaceae, Mycoblastaceae, Pilocarpaceae, Psoraceae, Ramalinaceae, Scoliciosporaceae, and Squamarinaceae.

The sister group relation of parmeliaceae (Lecanorales, Ascomycota).

The family Parmeliaceae (Lecanorales, Ascomycota) is possibly the largest, best known and most thoroughly studied lichen family within its order. Despite this fact the relationship between Parmeliaceae and other groups in Lecanorales is still poorly known. The aim of the present study is to contribute to finding the sister group of Parmeliaceae as an aid in future studies on the phylogeny and character evolution of the group. We do this by sampling all potential relatives to the Parmeliaceae that we have identified, i.e. Gypsoplaca, Japewia, Mycoblastus, Protoparmelia, and Tephromela, a good representation of the major groups within the Parmeliaceae s. lat. and a good representation of other taxa in the core Lecanorales. We use molecular data from two genes, the large subunit of the nuclear ribosomal RNA gene (nrLSU) and the small subunit of the mitochondrial ribosomal RNA gene (mrSSU), and a Bayesian analysis of the combined data. The results show that the closest relatives to Parmeliaceae are the two genera Protoparmelia and Gypsoplaca, which are crustose lichens. Parmeliaceae in our sense is a well supported group, including also the family segregates Alectoriaceae, Hypogymniaceae, Usneaceae and Anziaceae.

Molecular phylogeny of the genus Lecania (Ramalinaceae, lichenized Ascomycota).

The molecular phylogeny of the lichen genus Lecania was investigated using nucleotide sequences from the mt-SSU rRNA, the ITS region of the nu-rDNA, and the RNA polymerase II second largest subunit. Forty-six species representing Lecania and other genera likely to influence the phylogeny were included in the study. Phylogenetic reconstructions were carried out using Bayesian inference, ML, and MP approaches. Lecania, as traditionally circumscribed, is not a monophyletic genus. However, a monophyletic group containing a large number of Lecania species, including the type species L. fuscella, was discovered in the analysis, and recognition of Lecania sensu stricto is suggested. L. baeomma, L. glauca, L. gerlachei, L. brialmontii, L. racovitzae, L. hyalina (alias Biatora globulosa), L. chlorotiza, L. naegelii, and L. furfuracea do not belong in Lecania s. str., although the latter two are closely related to Lecania s. str. Representatives of the genus Bilimbia form a well-supported group, as does the 'Thamnolecania' group containing the Antarctic 'Lecania' species, L. gerlachei, L. brialmontii, and L. racovitzae. An alternative to recognizing these two genera would be a wider circumscription of Bilimbia to include the 'Thamnolecania' group as well as affiliated taxa.

Genetic variation and population differentiation in the lichen-forming ascomycete Xanthoria parietina on the island Storfosna, central Norway.

Genetic diversity and fine-scale population structure in the lichen-forming ascomycete Xanthoria parietina was investigated using sequence variation in part of the intergenic spacer (IGS) and the complete internal transcribed spacer (ITS) regions of the nuclear ribosomal DNA. Sampling included 213 and 225 individuals, respectively, from seven populations in two different habitats, bark and rock, on the island Storfosna off the central west coast of Norway. Both markers revealed significant variation and a total of 10 IGS and 16 ITS haplotypes were found. There were no signs of significant positive spatial autocorrelation at any spatial size class down to 10% of transect length, nor did we find significant deviations from neutrality or signs of historical population expansion. Analysis of molecular variance (amova) indicated that most of the genetic variance observed was within populations, but when populations were grouped according to habitat, more than a quarter of the variance was explained among groups. Pairwise comparisons of populations (F(ST), exact tests of population differentiation) revealed significant differentiation between populations in different habitats (on bark or rock), but not between populations in the same habitat. Haplotype networks show that internal and presumably old haplotypes are shared between habitats, whereas terminal haplotypes tend to be unique to a habitat, mostly bark. We interpret the observed pattern to mean that there is no evidence of restricted gene flow between populations in the same habitat at the present spatial scale (interpopulation distances one or a few kilometres). On the other hand, differentiation between habitats is considerable, which we attribute to restricted gene flow between habitats (habitat isolation). Evidence suggests that the observed differentiation did not evolve locally. Estimates of divergence time between populations in the respective habitats indicate that an ancestral population started to diverge at least 34,000 years ago but probably much further back in time.

Fungal diversity in rock beneath a crustose lichen as revealed by molecular markers.

Lichen-forming fungi have been assumed to be more or less restricted to the surface of the substrate on which they grow, Conclusive identification of hyphae or an assessment of the fungal diversity inside lichen-covered rock has not been possible using methods based on direct observation. We circumvented this problem by using a DNA sequencing approach. Cores were drilled from a Devonian arcosic sandstone rock harboring the crustose lichen Ophioparma ventosa (L.) Norman on the surface. The cores were cut vertically, and DNA was extracted from the pulverized rock slices. A series of polymerase chain reactions using fungal-specific primers as well as Ophioparma ventosa specific primers were employed to amplify the internal transcribed spacer region of the nuclear ribosomal DNA. The results show that hyphae of O. ventosa penetrate approximately 10-12 mm into the rock. Consequently, the hyphal layer formed by the lichen fungus inside the rock could be 7-12 times as thick as the symbiotic thallus at the surface of the rock. In addition, eight non-lichenized fungal taxa and five that could not be identified to species were encountered. One fungal species in the order Helotiales occurs in six of the eight cores. The significance of these results to the colonization and weathering of rock by lichenized fungi is discussed.

Phylogenetic relationships of Lecanoromycetes (Ascomycota) as revealed by analyses of mtSSU and nLSU rDNA sequence data.

The phylogeny of Lecanoromycetes (Ascomycota, Fungi) is investigated utilizing parsimony and Bayesian Markov Chain Monte Carlo analyses, of combined nLSU rDNA and mtSSU rDNA sequence datasets. The results suggest that Acarosporaceae, Candelariaceae, Phlyctis and Pycnora are not members of the monophyletic Lecanorales, and that Timdalia and Pleopsidium are members of a monophyletic Acarosporaceae. Pycnora, Candelariaceae and Acarosporaceae form a monophyletic group. Umbilicariaceae, Hypocenomyce scalaris, H. friesii, Ophioparmaceae, Boreoplaca, Elixia and Fuscidea form either a basal paraphyletic assemblage in Lecanoromycetes, or a monophyletic group which is the sister-group to Lecanorales and the rest of Lecanoromycetes (excluding Acarosporaceae). The Acarosporaceae forms a group with Pycnora and Candelariaceae, which may be outside the Lecanoromycetes. Chaetothyriales, Verrucariales, Eurotiales, Lichinales and Mycocaliciales form a monophyletic group, but with low support. We briefly discuss incongruence between datasets from different genetic markers, comparing the differences between the separate parsimony analyses, where the ILD test indicated a very significant incongruence. The phylogenetic significance of ascus-types that have influenced most recent Ascomycota classifications heavily is also discussed, and we finally point out risks with formalizing classifications too early.

Molecular evidence supports the distinction between Xanthoria parietina and X. aureola (Teloschistaceae, lichenized Ascomycota).

This study aims to clarify taxonomic relationships within the current concept of Xanthoria parietina in northern Europe. For comparison, X. calcicola was also included in the study. Morphological as well as molecular data were utilized. Morphology indicated the presence of three species, Xanthoria parietina, X. calcicola, and X. aureola, the latter of which is resurrected here from synonymy. The most important separating characters involve colour and thickness of the thallus, lobe width, morphology of laminar structures, and the texture of the upper surface. X. aureola, as recognized here, mostly occurs on seashore rocks. Part of the IGS region as well as the complete ITS were sequenced in 70 individual thalli representing ten geographical regions in Europe. In total, 19 different IGS haplotypes and 20 different ITS haplotypes were present in the data set. Owing to indications of possible recombination between the IGS and the ITS, the two data sets were analyzed separately. Haplotype networks were estimated, both of which indicate that X. parietina is distinct from X. aureola and X. calcicola. In our sample, the two latter do not share haplotypes, but are only separated by a few mutational steps.

Disintegration of the Micareaceae (lichenized Ascomycota): a molecular phylogeny based on mitochondrial rDNA sequences.

The phylogeny of the family Micareaceae and the genus Micarea was studied using mitochondrial small subunit ribosomal DNA sequences. Phylogenetic reconstructions were performed using Bayesian MCMC tree sampling and a maximum likelihood approach. The Micareaceae in its current sense is highly heterogeneous, and Helocarpon, Psilolechia, and Scutula, all thought to be close relatives of Micarea, are shown to be only distantly related. The genus Micarea is paraphyletic unless the entire Pilocarpaceae and Ectolechiaceae are included, as also indicated by an expected likelihood weights test. It is suggested that the Micareaceae is reduced to synonymy with the Pilocarpaceae, which also includes the Ectolechiaceae, and that Micarea may have to be divided into a series of smaller genera in the future. Micarea species with a 'non-micareoid' photobiont group with Psora and the Ramalinaceae, whereas Micarea intrusa appears to belong in Scoliciosporum. Three species fall inside the paraphyletic Micarea: Szczawinskia tsugae, Catillaria contristans, and Fellhaneropsis vezdae. Tropical foliicolous taxa are nested within groups of mainly temperate and arctic-alpine distribution. A 'micareoid' photobiont appears to be plesiomorphic in the Pilocarpaceae but has been lost a few times.

Evolution and taxonomy of the marine Collemopsidium species (lichenized Ascomycota) in north-west Europe.

The taxonomy of the marine species of Collemopsidium in northwest Europe was investigated using morphological and molecular evidence. 210 specimens were collected from the west coasts of Norway and Ireland, and morphological and ecological variables recorded. ITS1 rDNA sequences were obtained from 24 specimens. A phylogenetic analysis, resulting in a single optimal tree, was performed under the unweighted least squares optimality criterion based on maximum likelihood distances obtained from unaligned sequences. Principal components analysis (PCA) was performed on morphological variables of the sequenced specimens, and classification was carried out by maximizing agreement between the phylogenetic tree and the PCA. Thallus immersion, and perithecial immersion and size, were the most important characters for discriminating between taxa. Apart from substratum, niche separation between taxa was small but statistically significant as shown by a redundancy analysis (RDA). Variance partitioning indicated that genetic variation is vastly more important than ecology for explaining phenotypic variation. Five species of marine Collemopsidium are recognized, including two new combinations: C. foveolatum (syn. Arthopyrenia foveolata) and C. ostrearum (syn. Lecanactis ostrearum) combs. nov.

Supraordinal phylogenetic relationships of Lecanoromycetes based on a Bayesian analysis of combined nuclear and mitochondrial sequences.

Phylogenetic relationships of lichen-forming discomycetes and their relatives in the class Lecanoromycetes were examined by using nuclear large subunit and mitochondrial small subunit ribosomal DNA sequences. Ninety-eight partial sequences of 53 ascomycetes were generated and aligned with the corresponding sequences retrieved from GenBank resulting in an alignment of 100 taxa that was analyzed using a Bayesian approach with Markov chain Monte Carlo (B/MCMC) methods. The analysis revealed the monophyly of the Lecanoromycetes with two major clades: one clade including the monophyletic orders Graphidales and Ostropales and the paraphyletic Gyalectales, the other clade including the monophyletic Lecanorales (incl. Caliciales, Peltigerales, and Teloschistales) and a clade containing the polyphyletic Agyriales, a yet undescribed order Umbilicariales (including Elixiaceae and Umbilicariaceae), and Pertusariales. The monophyly of the Pertusariales was not resolved. Testing of alternative hypotheses revealed that a placement of Chaetothyriomycetes and Eurotiomycetes within Lecanoromycetes and the monophyly of Agyriales s. lat. (incl. Elixiaceae and Schaereriaceae) and Ostropales s. lat. (incl. Graphidales) can be rejected, while monophyly of Gyalectales and the Pertusariales and placement of Umbilicariales on the Lecanorales branch cannot be rejected with the current data set.

Local population subdivision in the lichen Cladonia subcervicornis as revealed by mitochondrial cytochrome oxidase subunit 1 intron sequences.

A 753-771 bp long intronic sequence from the mitochondrial cox 1 gene of Cladonia subcervicornis (Cladoniaceae, Lecanorales, Ascomycota) was amplified with newly designed PCR primers. The cox 1 intron sequence, which apparently has not been used for phylogenetic or population genetic research in fungi, displays high infraspecific variation. Sequences were obtained from 124 specimens from four neighboring localities in coastal Hordaland, western Norway. An exact test of population differentiation and population pairwise fixation indices F(ST) show significantly reduced gene flow between the northernmost locality and the other three populations. Although Cladonia subcervicornis frequently produces apothecia, we conclude that dispersal by ascospores over long distances is rather ineffective in this species.

Tree-Dependent Lichens and Beetles as Indicators in Conservation Forests.

We tested whether the conspicuous lichen Lobaria pulmonaria indicates the number of tree-dependent, red-listed species in a hemiboreal forest in southern Sweden. In 18 naturally regenerated, mainly old deciduous forest plots considered to be of high or very high conservation value, the number of red-listed tree lichens or wood beetles was not positively correlated with the area of the forest stands studied (8-56 ha). The 8 stands with L. pulmonaria had about nine (median) red-listed lichens, but 10 stands without L. pulmonaria had only about one such species, a highly significant difference. Similarly, the variation between stands in the number of red-listed wood beetles was considerable, but it was very weakly correlated with the number of red-listed lichens. The number of red-listed wood beetles dependent on dead trees was not different in areas with or without Lobaria pulmonaria, but the number of red-listed wood beetles dependent on hollow trees was higher in stands with Lobaria pulmonaria (median of seven species) than in those without (three species) the lichen. Stands with this lichen species also contained significantly more of other types of lichens that have been proposed as indicators of forest continuity according to three lists. We stress the need for identification of species that could serve as indicators of different types of forest continuity and identify some organism groups that may indicate the different types. Los líquenes y coleópteros dependientes de árboles como indicadores en los bosques bajo conservación.

Resumen: Evaluamos si el consipicuo líquen Lobaria pulmonaria indica el número de especies dependientes de los árboles, listadas en el libro rojo en un área de bosque hemi-boreal en el sur de Suecia. El número de líquenes de los árboles o coleópteros de la madera listados en el libro rojo no estuvo positivamente correlacionado con el área de los rodales de bosque estudiados (8-56 ha), en 18 rodales naturalemente regenerados, en su mayoría de antiguo crecimiento de bosques deciduos; los cuáles se considera tienen de alto a muy alto valor de conservación. Los 8 rodales con L. pulmonaria tuvieron alrededor de nueve (mediana) líquenes listados en el libro rojo, pero 10 rodales sin L. pulmonaria tuvieron sólamente una de dichas especies, una diferencia significativamente alta. Asi mismo, la variación entre rodales en el número de coleópteros listados en el libro rojo fue considerable, pero estuvo debilmente correlacionada con el número de líquenes listados en el libro rojo. El número de coleópteros de la madera listados en el libro rojo, dependientes de árboles muertos no fue diferente en áreas con o sin Lobaria pulmonaria. Sin embargo, el número de coleópteros de la madera listados en el libro rojo dependientes de troncos huecos fue más alto en rodales con Lobaria pulmonaria (mediana = siete especies) que en aquellos sin el mencionado líquen (tres epecies). Los rodales con este líquen también contuvieron significativamente un número mayor de otros líquenes que han sido propuestos como indicadores de la continuidad del bosque de acuerdo a tres listas. Remarcamos la necesidad de la identificación de especies que pueden servir como indicadoras de los diferentes tipos de continuidad del bosque y la identificación de ciertos grupos de organismos que pueden indicar los diferentes tipos.