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Members of the poorly investigated family Teloschistaceae in South America, mostly from Bolivia and Peru, were examined using molecular and morphological data here for the first time. In recent phylogenetic reclassifications of Teloschistaceae, South American representatives were poorly represented but shown to belong to subfamilies Teloschistoideae and Xanthorioideae. In this study, we expanded the sampling of South American taxa and investigated mainly the lobate, sublobate, and squamulose members of Caloplaca s.l., using morphological characters and a molecular phylogeny based on a combined three-locus data set (one mitochondrial and two nuclear loci). Building upon new phylogenies at the family and subfamily levels (Teloschistoideae), we propose here three new genera: Andina, Aridoplaca , and Cinnabaria , with the type species Andina citrinoides, Aridoplaca peltata , and Cinnabaria boliviana . We also propose to reduce Tarasginia to synonymy with Sirenophila and Tayloriellina to synonymy with Villophora and introduce three new combinations: Dufourea ottolangei, D. volkmarwirthii , and Villophora erythrosticta. Scutaria andina is reported as new to Bolivia. A critical revision of the subfamily Brownlielloideae confirmed recent findings that it is an artifactual taxon based on a "chimeric" data set, with the type genus being part of Teloschistoideae.

Foliicolous lichens are formed by diverse, highly specialized fungi that establish themselves and complete their life cycle within the brief duration of their leaf substratum. Over half of these lichen-forming fungi are members of either the Gomphillaceae or Pilocarpaceae, and associate with Trebouxia-like green algae whose identities have never been positively determined. We investigated the phylogenetic affinities of these photobionts to better understand their role in lichen establishment on an ephemeral surface. Thallus samples of Gomphillaceae and Pilocarpaceae were collected from foliicolous communities in southwest Florida and processed for sequencing of photobiont marker genes, algal cultivation and/or TEM. Additional specimens from these families and also from Aspidothelium (Thelenellaceae) were collected from a variety of substrates globally. Sequences from rbcL and nuSSU regions were obtained and subjected to Maximum Likelihood and Bayesian analyses. Analysis of 37 rbcL and 7 nuSSU algal sequences placed all photobionts studied within the provisional trebouxiophycean assemblage known as the Watanabea clade. All but three of the sequences showed affinities within Heveochlorella, a genus recently described from tree trunks in East Asia. The photobiont chloroplast showed multiple thylakoid stacks penetrating the pyrenoid centripetally as tubules lined with pyrenoglobuli, similar to the two described species of Heveochlorella. We conclude that Heveochlorella includes algae of potentially major importance as lichen photobionts, particularly within (but not limited to) foliicolous communities in tropical and subtropical regions worldwide. The ease with which they may be cultivated on minimal media suggests their potential to thrive free-living as well as in lichen symbiosis., (© 2016 Phycological Society of America.)

The Agaricales is the largest and most diverse order of mushroom-forming Basidiomycota, with over 100 natural groups recognized in recent Fungal Tree of Life studies. Most agarics are either saprotrophic or ectomycorrhizal fungi, but the family Hygrophoraceae is in part characterized by a unique and remarkable diversity of lichenized forms. The most familiar of these is the chlorolichen genus Lichenomphalia, whose phylogenetic position in the Agaricales has been established. Recent limited evidence suggested that Hygrophoraceae also contains cyanolichens in the genus Dictyonema, which indicates a remarkable concentration and diversity of lichen-formers in a single family of agarics. To demonstrate the relationships of lichen-formers to other fungi in the family, we assembled ribosomal sequences from 52 species representing recognized groups within the Hygrophoraceae, among them new sequences representing Acantholichen and most species and forms of Dictyonema. The molecular data were evaluated using parsimony, likelihood, Bayesian, and distance analyses, including coding of ambiguous regions by means of INAASE and ARC, all of which indicate that Dictyonema and Acantholichen form a monophyletic clade derived from the primarily bryophilous genus Arrhenia and sister to the enigmatic Athelia pyriformis, a species unrelated to the Atheliales for which we are proposing a new genus name Eonema. The chlorolichen genus Lichenomphalia may be polyphyletic. Fungi in the Dictyonema-Acantholichen clade are typically tropical, entirely lichenized, and associate with cyanobacterial photobionts. Our data indicate a transition from agaricoid-omphalinoid basidiomes observed in Arrhenia to stereoid-corticioid forms in Dictyonema, and also support a previous suggestion of a connection between loss of clamp connections and lichenization. The diverse basidiome and thallus morphologies and nutritional ecologies of these fungi indicate a remarkable evolutionary flexibility that appears to have developed in part as a consequence of symbiosis.

The foliicolous lichen Gallaicolichen pacificus exhibits unique goniocystangia-like structures named peltidiangia and peltidia. Its taxonomic classification within the Ascomycota has been unclear due to the absence of ascomata and lack of molecular data. Here we clarify the phylogenetic affinities of Gallaicolichen pacificus by analyzing mitochondrial small subunit ribosomal RNA (mtSSU) sequences obtained from specimens collected in New Caledonia. Ascomata and ascospores of G. pacificus , previously unknown, are described and illustrated for the first time. The results from the molecular and morphological analyses clearly indicate that Gallaicolichen pacificus belongs to the Porinaceae and is closely related to Porina guianensis. This is a remarkable extension of the already known, wide morphological diversity of thalli and diaspores produced within this family. [ABSTRACT FROM AUTHOR]

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The genus Dendriscosticta (Ascomycota: Peltigerales) encompasses several distinctive lichen‐forming fungal species restricted to the Northern Hemisphere. Most are flagship species of old‐growth forests with good air quality. A global phylogeny of the genus based on multilocus sequence data (ITS, RPB1, EF‐1α, MCM7), model‐based phylogenetic methods, and morphological and chemical assessments, reveals a high level of cryptic speciation often associated with restricted geographical distribution and/or chemical characters. Using sequence‐based species delimitation approaches, we circumscribe two main clades referred to as the D. wrightii clade, with five unequivocal species, including D. gelida sp. nov., and the D. praetextata clade, with eight putative species, including D. phyllidiata sp. nov. The absence of recently collected material of D. hookeri comb. nov. from the type locality unfortunately prevents assignment of this epithet to one of the five supported lineages sharing this morphotype. Three new combinations are proposed: D. hookeri, D. insinuans comb. nov. and D. yatabeana comb. nov. Epitypes are designated for D. wrightii and D. yatabeana. Species diversity within the genus increased from four to nine. Our morphological assessment confirmed that Sticta and Dendriscosticta can be readily distinguished by the presence of excipular algae whereas the structure of the lower surface pores is not a reliable diagnostic feature. [ABSTRACT FROM AUTHOR]

Three species of lichenized basidiomycetes in the Dictyonema clade from southeastern North America are described as new to science: Cyphellostereum georgianum, C. jamesianum and Dictyonema lawreyi, all with a crustose-filamentous growth form. Based on ITS sequences, the species form well-supported monophyletic clades in a phy-logeny and are represented by at least two specimens each. They are also distinguishable by morphological and anatomical characters. These new findings emphasize the importance of lichenological studies in North America, especially in historically understudied taxonomic groups, such as basidiolichens. This study is dedicated to James D. Lawrey on the occasion of his 70th birthday. [ABSTRACT FROM AUTHOR]

Phylogenetic studies revealed a high level of diversity within the lichen-forming fungus so far identified with the name Maronina multifera (Protoparmelioideae, Parmeliaceae), currently accommodated in the genus Neoprotoparmelia. Here, six new species of Neoprotoparmelia are described as new to science based on morphological and molecular data, mostly from northeastern Brazil. The new species are: Neoprotoparmelia nigra (Brazil), with 32-spored asci, lacking alectoronic acid, and with pale, K-negative apothecial base, and blackish apothecial disc; N. paramultifera (Brazil), with 64-spored asci, alectoronic acid, pale, K-negative apothecial base, and purplish brown apothecial disc and thick apothecial margins; N. pseudomultifera (Brazil), with 32-spored asci, lacking alectoronic acid, and with pale, K-negative apothecial base and brown apothecial disc (no reddish or purplish tinge); N. purpurea (Brazil), with 32-spored asci, lacking alectoronic acid, and with pigmented, K+ purplish-violet apothecial base and purplish brown apothecial disc; N. rubrofusca (Colombia), with 32-spored asci, lacking alectoronic acid, and with pigmented, K+ purplish-violet apothecial base, and red-brown apothecial disc and thin, evanescent margins; and N. sexdecimspora (Brazil), with 16-spored asci, alectoronic acid, pale, K-negative apothecial base, and purplish brown apothecial disc. The name N. multifera is restricted to a species from the northern Andes with 64-spored asci, alectoronic acid, pale, K-negative apothecial base, and purplish brown apothecial disc with thin margins, while the new combination N. camptotheca is adopted for a species in eastern Brazil with 32-spored asci, alectoronic acid, pale, K-negative apothecial base, purplish brown apothecial disc, and smooth margin (all other species in the complex having crenulate margins). The following two new combinations are also proposed: Neoprotoparmelia saxicola and N. rogersii (syn.: N. capensis V.J.Rico, A.Crespo & Garima Singh). [ABSTRACT FROM AUTHOR]

Heterocyphelium is a mazaediate genus containing a single species, H. leucampyx. The species was originally described from Cuba within the genus Trachylia (Arthoniales, Arthoniaceae) and later placed in various genera of the collective order Caliciales s. lat. For the past three decades, Heterocyphelium was considered an orphaned genus (incertae sedis) within the Ascomycota, since morphology alone could not resolve its systematic position. In this study, we added molecular data with the aim of resolving this uncertainty. Bayesian and maximum likelihood analyses of newly generated sequence data from the mitochondrial ribosomal RNA small subunit (mtSSU) and the RNA polymerase II second largest subunit gene (RPB2) provide clear evidence that Heterocyphelium leucampyx is nested within the order Arthoniales, in the family Lecanographaceae, sister to the genus Alyxoria. Heterocyphelium is a further example of parallel evolution of passive spore dispersal, prototunicate asci and the occurrence of a mazaedium in the Ascomycota, and another calicioid genus whose systematic placement could be eventually clarified by means of molecular data. Heterocyphelium is the fourth mazaediate genus in Arthoniales, in addition to Sporostigma, Tylophorella and Tylophoron. [ABSTRACT FROM AUTHOR]

As part of a phylogenetic and taxonomic revision of the lichenized genus Sticta in Colombia, six new species with apotheciate thalli but lacking vegetative propagules are described: Sticta atroandensis Moncada & Lücking sp. nov., S. brevior Moncada & Lücking sp. nov., S. lumbschiana Moncada & Lücking sp. nov., S. macrocyphellata Moncada & Coca sp. nov., S. parahumboldtii Moncada & Lücking sp. nov., and S. pseudohumboldtii Moncada & Lücking sp. nov. The last two are related to S. humboldtii but phylogenetically distinct and differ in details of cyphellae morphology and spot reactions. Sticta atroandensis is similar to S. andensis but has a dark lower tomentum, whereas S. brevior belongs in the S. kunthii group, being characterized by mainly laminal apothecia with tomentose margins and a rather stout thallus. Both S. lumbschiana and S. macrocyphellata form part of the S. dilatata group and can be characterized by their lobe surface structure and cyphellae anatomy. Keys to species of the S. dilatata and S. humboldtii groups are presented. [ABSTRACT FROM PUBLISHER]

We studied ascospore dispersal and ontogeny in several species of Trypetheliaceae and one species of Graphidaceae with megalosporous (very large) ascospores, including Aptrootia terricola (Aptroot) Lücking, Umaña & Chaves, Architrypethelium nitens (Fée) Aptroot, A. seminudum (Mont.) Aptroot, Astrothelium diplocarpoides Müll. Arg., Laurera gigantospora (Müll. Arg.) Zahlbr., and Ocellularia subpraestans Hale. Although mature ascospores are very different among species of Trypetheliaceae, all species studied share plesiomorphic ontogenetic features such as the formation of a single central euseptum and subsequent development of a variable number of transverse distosepta with diamond-shaped lumina (astrothelioid ascospore type). Even the dark brown, richly muriform ascospores of Aptrootia terricola go through an astrothelioid stage. This suggests that ascospore types may be more closely related than suggested by their mature morphology, and could explain why related taxa can develop markedly different ascospore types. We discuss the implications for systematic classification of Trypetheliaceae in the light of recent molecular studies, and also speculate about the ecological importance of large ascospores, especially with muriform septation. [ABSTRACT FROM AUTHOR]

Phylogenetic diversity of lichen photobionts is low compared to that of fungal counterparts. Most lichen fungi are thought to be associated with just four photobiont genera, among them the cyanobacteria Nosioc and Scytoneina, two of the most important nitrogen fixers in humid ecosystems. Although many Nostoc photobionts have been identified using isolated cultures and sequences, the identity of Scytoneina photobionts has never been confirmed by culturing or sequencing. We investigated the phylogenetic placement of presumed Scytonema photobionts and unicellular morphotypes previously assigned to Chroococcus, from tropical Dictyonerna, Acantholichen, Coccocarpia, and Stereocaulon lichens. While we confirm that filamentous and unicellular photobiont morphotypes belong to a single dade, this dade does not cluster with Scytonema but represents a novel, previously unrecognized, highly diverse, exclusively lichenized lineage, for which the name Rhizonerna is available. The phylogenetic structure observed in this novel lineage suggests absence of coevolution with associated mycobionts at the species or dade level. Instead, highly efficient photobiont strains appear to have evolved through photobiont sharing between unrelated, but ecologically similar, coexisting lineages of lichenized fungi ("lichen guilds"), via the selection of particular photobiont strains through and subsequent horizontal transfer among unrelated mycobionts, a phenomenon not unlike crop domestication. [ABSTRACT FROM AUTHOR]

Aim To analyse the historical biogeography of the lichen genus Chroodiscus using a phenotype-based phylogeny in the context of continental drift and evolution of tropical rain forest vegetation. Location All tropical regions (Central and South America, Africa, India, Southeast Asia, north-east Australia). Methods We performed a phenotype-based phylogenetic analysis and ancestral character state reconstruction of 14 species of the lichen genus Chroodiscus, usingpaup* andmesquite; dispersal–vicariance analysis (DIVA) and dispersal–extinction–cladogenesis (DEC) modelling to trace the geographical origin of individual clades; and ordination and clustering by means ofpc-ord, based on a novel similarity index, to visualize the biogeographical relationships of floristic regions in which Chroodiscus occurs. Results The 14 species of Chroodiscus show distinctive distribution patterns, with one pantropical and one amphi-Pacific taxon and 12 species each restricted to a single continent. The genus comprises four clades. DIVA and DEC modelling suggest a South American origin of Chroodiscus in the mid to late Cretaceous (120–100 Ma), with subsequent expansion through a South American–African–Indian–Southeast Asian–Australian dispersal route and late diversification of the argillaceus clade in Southeast Asia. Based on the abundance of extant taxa, the probability of speciation events in Chroodiscus is shown to be extremely low. Slow dispersal of foliicolous rain forest understorey lichens is consistent with estimated phylogenetic ages of individual species and with average lengths of biological species intervals in fungi (10–20 Myr). Main conclusions The present-day distribution of Chroodiscus can be explained by vicariance and mid-distance dispersal through the interconnection or proximity of continental shelves, without the need for recent, trans-oceanic long-distance dispersal. Phylogenetic reconstruction and age estimation for Chroodiscus are consistent with the ‘biotic ferry’ hypothesis: a South American origin and subsequent eastward expansion through Africa towards Southeast Asia and north-eastern Australia via the Indian subcontinent. The present-day pantropical distributions of many clades and species of foliicolous lichens might thus be explained by eastward expansion through continental drift, along with the evolution of modern rain forests starting 120 Ma, rather than by the existence of a hypothetical continuous area of pre-modern rain forest spanning South America, Africa and Southeast Asia during the mid and late Cretaceous. [ABSTRACT FROM AUTHOR]

Abstract: Phylogenetic studies indicate that the basidiolichen genus Dictyonema s.lat., often thought to represent only a single genus with few species, includes several well-supported genus-level clades, all of which form associations with a unique lineage of obligately lichenized cyanobacteria (Rhizonema). In an attempt to elucidate the evolution and genus- and species-level diversification in Dictyonema s.lat., we generated 68 new sequences of the nuclear large subunit rDNA (nuLSU), the internal transcribed spacer (ITS), and the RNA polymerase II subunit (RPB2), for 29 species-level lineages representing all major clades of Dictyonema s.lat. and most of the species currently known. The multilocus phylogeny obtained via maximum likelihood and Bayesian approaches indicates the presence of five genus-level groups: a basal clade, Cyphellostereum, that is sister to the rest of the species, a paraphyletic grade representing Dictyonema s.str., and three clades representing the genera Acantholichen, Cora, and Corella. To determine the evolutionary transformations of the lichenized thallus in the group, ancestral character state reconstruction was done using six characters (lichenisation, thallus type, cortex type, hyphal sheath and haustorial type, photobiont morphology, and basidiocarp type). Our analysis indicates a progressive development of the lichenized thallus from loosely organized filamentous crusts with separate, cyphelloid basidiocarps in Cyphellostereum, to filamentous crusts with derived hyphal sheath and cyphelloid–stereoid basidiocarps partially incorporated into the lichen thallus in Dictyonema, to squamulose–foliose thalli with corticioid basidiocarps entirely supported by the lichen thallus in Cora. These results indicate a remarkable evolutionary integration of lichenized and reproductive tissues in Dictyonema s.lat., supporting the hypothesis that, at least in this case, lichenized thalli may have evolved from reproductive structures in their nonlichenized ancestors. [Copyright &y& Elsevier]

Abstract: 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. [Copyright &y& Elsevier]

Abstract: A comprehensive phylogenetic classification of the kingdom Fungi is proposed, with reference to recent molecular phylogenetic analyses, and with input from diverse members of the fungal taxonomic community. The classification includes 195 taxa, down to the level of order, of which 16 are described or validated here: Dikarya subkingdom nov.; Chytridiomycota, Neocallimastigomycota phyla nov.; Monoblepharidomycetes, Neocallimastigomycetes class. nov.; Eurotiomycetidae, Lecanoromycetidae, Mycocaliciomycetidae subclass. nov.; Acarosporales, Corticiales, Baeomycetales, Candelariales, Gloeophyllales, Melanosporales, Trechisporales, Umbilicariales ords. nov. The clade containing Ascomycota and Basidiomycota is classified as subkingdom Dikarya, reflecting the putative synapomorphy of dikaryotic hyphae. The most dramatic shifts in the classification relative to previous works concern the groups that have traditionally been included in the Chytridiomycota and Zygomycota. The Chytridiomycota is retained in a restricted sense, with Blastocladiomycota and Neocallimastigomycota representing segregate phyla of flagellated Fungi. Taxa traditionally placed in Zygomycota are distributed among Glomeromycota and several subphyla incertae sedis, including Mucoromycotina, Entomophthoromycotina, Kickxellomycotina, and Zoopagomycotina. Microsporidia are included in the Fungi, but no further subdivision of the group is proposed. Several genera of ‘basal’Fungi of uncertain position are not placed in any higher taxa, including Basidiobolus, Caulochytrium, Olpidium, and Rozella. [Copyright &y& Elsevier]

Abstract: The phylogenetic position of Trypetheliaceae was studied using partial sequences of the mtSSU and nuLSU rDNA of 100 and 110 ascomycetes, respectively, including 48 newly obtained sequences. Our analysis confirms Trypetheliaceae as monophyletic and places the family in Dothideomycetes. Pyrenulaceae, which were previously classified with Trypetheliaceae in Pyrenulales or Melanommatales, are supported as belonging to Chaetothyriomycetes. Monophyly of Pyrenulales, including Trypetheliaceae is rejected using three independent test methods. Monophyly of Arthopyreniaceae plus Trypetheliaceae, the two families including lichen-forming fungi in Dothideomycetes, is also rejected, as well as a placement of Trypetheliaceae in Pleosporales (incl. Melanommatales). [Copyright &y& Elsevier]