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.

Most lichen-forming fungi are characterized by the production of secondary metabolites. Differences in metabolite patterns have frequently served to distinguish lichen taxa with subsequent controversies about the rank of chemical variants (chemotype, variety, subspecies or species). Using a model system, we investigate whether production of norstictic acid within a group of lichenized ascomycetes is correlated with phylogenetic patterns, population differentiation or single and multi-locus haplotypes. Our study is based on DNA sequences of three gene loci (ITS, GPD, mtLSU) together with HPLC (311) and TLC (594) data from a total of 594 samples of three closely related fruticose lichens: Cetraria aculeata and C. muricata without norstictic acid, and C. steppae with norstictic acid. In nature, C. aculeata and C. steppae often occur together and the status of C. steppae as a separate species has been questioned. Our results show geographical but no phylogenetic structure of norstictic acid production and few significant associations between genetic clusters and the occurrence of norstictic acid. All frequently distributed haplotypes display differences in norstictic acid content. The few associations at the population level are most likely a by-product of spatial genetic structure, because norstictic acid was expressed only in individuals from the Mediterranean-Central Asian part of the study area. We conclude that the production of norstictic acid in the C. aculeata group is most likely triggered by the environment (climate, edaphic factors, associated symbionts). Cetraria steppae might be a different evolutionary lineage restricted to warm temperate regions but it is not uniquely characterized by the presence of norstictic acid.

The crustose lichen biota of coastal rocks in South Brazil was investigated. Several distinct lichen zones were found: a littoral black zone, a supralittoral yellow zone and a grey zone with species restricted to either exposed granite, dry overhangs, damp overhangs or places subjected to run-off. Dendrographa austrosorediata is newly described, with a crustose, flat to partly curling up or blister-like thallus which is much dissected, whitish grey, 0·1–0·2 mm thick; surface minutely densely cracked, rimose, with cracks intersecting at each c. 0·1 mm; soredia whitish to bluish grey but asymmetrically blackened in the direction facing the light, originating on the thallus surface, in initially discrete convex soralia. The phylogenetic position of this new species was traced by molecular methods. Stigmidium marinum, generally regarded as a lichenicolous fungus, was found as a free-living lichen, thousands of kilometres away from the nearest known occurrence of any purported host.

Biological soil crusts are communities composed of cryptogamic organisms such as lichens, mosses, cyanobacteria and green algae that form a skin on soils in areas where vascular plants are excluded or limited by water availability or temperature. The lichen Psora decipiens (Hedw.) Hoffm. is a characteristic key organism in these communities in many different biomes. The species has a generalistic ecology and high morphological variation, which contributes to the ability of the species to withstand environmental changes. We investigated whether different populations, based on site and associated morpho-anatomical differences, incorporate functional water relations and how/whether this was driven by changes in abiotic factors. Samples were collected from two climatically distinct sites, one ‘dry’ site in southern Spain and one ‘wet’ site in the Austrian Alps. Our results showed that samples from the dry site had a significantly thicker epinecral layer, higher specific thallus area, a faster water uptake and contained more water per dry mass, all of which contributed to a much slower drying rate. Both populations showed a highly adjusted water gain that incorporates functional water relations and diffusion properties as a result of local water availability. We show eco-physiological and morphological mechanisms that underlie the high variability in P. decipiens and suggest how these might provide ecological benefits for this generalist lichen species.

Photosynthetic performance in lichens can vary throughout the year. We investigate the variation in the PSII quantum efficiency as a proxy for the physiological state of the photosynthetic apparatus in two umbilicate species from the genus Lasallia. Temporal variation in F v/F m in both species was monitored at a field site in Central Spain where both species coexist. Subsequent measurements were carried out in the laboratory after 48 h preconditioning. Both species showed clear variation during the year in PSII performance, with a marked depression in F v/F m during the summer. Lasallia pustulata consistently had higher F v/F m values than L. hispanica. Both species reached higher F v/F m values after 48 h of preconditioning in the laboratory and this recovery was particularly notable in the summer months. F v/F m was highly related to antecedent weather conditions during the two days prior to measurement.

This study aims to quantify the size-dependency of important hydration traits in Lobaria amplissima, L. pulmonaria and L. virens sampled in sympatric populations on deciduous tree trunks in southern Norway, and to discuss possible implications of species-specific traits for the ecological niches of these old forest cephalolichens. Traits measured were thallus size (area and mass), specific thallus mass (STM), internal (WHCinternal) and external water-holding capacity (WHCexternal), and water content (WC) after shaking and after blotting. Lobaria amplissima had the highest WHCinternal, 2·6 times higher than L. pulmonaria with the lowest WHCinternal. WHCinternal, driven by STM, strongly depended on size. WHCexternal was 28% (L. virens) to 47% (L. pulmonaria) of the WHCinternal. Unlike WHCinternal, WHCexternal did not depend on thallus area, meaning that WHCexternal is proportionally higher for smaller compared with larger thalli. The most widespread species, L. pulmonaria, benefits from a flexible hydration strategy due to low STM, facilitating the use of more diverse water sources than the other two species that depend more on rain, particularly L. amplissima with the highest STM and thus relatively high WHCinternal. For L. virens, a combination of less specialized hydration traits and a low tolerance to higher light intensity probably jeopardizes its survival outside rainforest habitats.

Numerous recent studies of lichenized fungi have uncovered hidden genetic diversity within a single phenotypic entity (so-called ‘cryptic species’). Here we report the opposite situation with vastly different morphologies apparently deriving from the same genotype. Endocena is a monotypic genus known only from southern South America. The single reported species, the terricolous E. informis, is morphologically variable; the type and other collections from the west coast of Chile being subfruticose, whereas specimens from further south and east are almost crustose in form. A sorediate terricolous lichen that is frequent on the Falkland Islands was confirmed by phylogenetic analysis of ITS rDNA and mtSSU rDNA sequences as being congeneric with E. informis and, surprisingly, both taxa were recovered as congeneric with the recently described genus and species Chirleja buckii, which is morphologically distinct from both E. informis and the sorediate taxon. Consequently, the genus Chirleja is included in the synonymy of Endocena and the new combination Endocena buckii is proposed. Because E. informis and the sorediate specimens have a similar thallus structure that differs radically from that of E. buckii, the name E. informis var. falklandica is proposed for the sorediate taxon. Poorly developed, incipient apothecia are also described from both varieties of E. informis, the first time that these have been reported for Endocena. We also report two lichenicolous fungi from E. informis var. informis, which are the first reports of lichenicolous fungi occurring on this genus.

The new lichenized ascomycete Tapellaria palaeotropica is described from Mahé Island in the Seychelles. The species is characterized by a crustose, pale green, smooth thallus dispersed in patches and by having black, rounded apothecia with flat discs, hyaline, transversely, 3–5-septate ascospores and a purplish brown excipulum. Morphology, distribution and related species are discussed. A world key to all currently known species in the genus is presented.

The new species Austrella isidioidea, which is unique in the genus in having isidioid structures on the thallus lobe ends as well as apothecia lacking a thalline margin, is described from the Falkland Islands. A collection with an identical mtSSU rDNA sequence to A. arachnoidea but with significant morphological differences (viz. a variable apothecial margin ranging from an arachnoid hyphal weft to a corticated regular margin) is reported from Îles Kerguelen. The phylogenetic and biogeographical implications of these new records are discussed.

Recent field trips to Amapá and the adjacent states of Amazonas and Pará uncovered unknown lichen species with unexpected relationships. The following new species of Arthoniales are described, mainly from Amapá in North Brazil: Neosergipea hyphosa, a corticolous species with an olive-green thallus, central part verrucose, marginal zone of 5–10 mm wide consisting of procumbent branching rhizohyphae, surrounded by a pinkish brown hyphal prothallus c. 3 mm wide into which the rhizohyphae protrude, globose ascomata, c. 0·1 mm diam., arranged in stromatoid groups of 5–15, with hyaline walls, without ascospores; Neosergipea septoconidiata, a corticolous species with a bluish grey thallus and bright orange, mostly conical pycnidia with fluffy hyphal surface, hyaline conidia, 3–6-septate, filiform, 12–23×1·5–2·0 μm, with rounded ends; Nyungwea pycnidiata, from a termite nest on a tree, with abundant pycnidia that are whitish, papillate, smooth, c. 0·1–0·2 mm diam., c. 0·2–0·4 mm high, at the base covered by thallus, and hyaline conidia that are simple, ellipsoid, 2·5–4·0×1·5–2·0 μm; and Opegrapha ramisorediata, a corticolous species with a thallus with initially rounded soralia that soon cover the complete thallus with a layer of fine yellowish green branched soredia, and containing gyrophoric acid. Apothecia are not known from any of these species but sequencing elucidated their generic affiliation. The genera Neosergipea and Nyungwea were previously monotypic and Nyungwea was known only from Africa. Nyungwea anguinella (Nyl.) Aptroot comb. nov. is a further new species in the latter genus, and through these two extra species we now know about the morphology of the pycnidia and apothecia of Nyungwea, which were unknown in the type species (a sterile crust).

The lowland area of southern Vietnam contains a high diversity of corticolous, crustose lichens, particularly in the family Graphidaceae. Two species, Acanthothecis verrucosa S. Joshi, Upreti & Hur and Graphis exuta S. Joshi, Upreti & Hur, are described new to science. Acanthothecis verrucosa is characterized by a verrucose, olive green thallus containing irregular to shortly lirellate apothecia, hyaline to brownish proper exciple apically lined by robust periphysoids, muriform ascospores, 30–70 × 15–20 μm, and the presence of psoromic and subpsoromic acids. Graphis exuta is characterized by a smooth, greenish white thallus, simple lirellae lacking thalline margin, entire labia coarsely white pruinose towards slit, muriform ascospores, 20–35×10–13 μm, and norstictic acid in the thallus. A new name, Acanthothecis yokdonensis S. Joshi & Hur, is proposed here for the species Acanthothecis salazinica S. Joshi & Hur, a later homonym of A. salazinica van den Boom & Sipman. In addition, 34 species from Vietnam are newly reported. A world key to the group of Graphis species characterized by short, simple and sessile lirellae lacking or with basal thalline margins (nuda-morph) is presented.

The lichen genus Ochrolechia found in China is revised. Twenty taxa are accepted in the genus, including four described as new to science: Ochrolechia alticola, differing from O. mexicana by the larger ascospores and the absence of lichexanthone, and from O. africana by the C+ red reaction of the apothecial margin cortex, the absence of 5-O-methylhiascic acid, and its alpine habitat; O. lijiangensis, differing from O. longispora by the larger apothecia and smaller ascospores, and by the presence of an additional undetermined fatty acid; O. longispora, differing from O. trochophora by the larger ascospores and the presence of two fatty acids; and O. rugomarginata, differing from O. pallescens in the minutely verruculose margins of the mature apothecia, the absence of variolaric acid and the C+ red thallus cortex, and from O. trochophora var. trochophora by the presence of alectoronic acid in the apothecial medulla, the coarsely pruinose disc and a preference for the bark of coniferous trees. Four new taxa are reported for China: Ochrolechia arborea, O. mahluensis, O. szatalaënsis and O. trochophora var. pruinirosella. An identification key to all taxa in the genus known from China is provided.

Rinodina is a widespread, polyphyletic genus of crustose Physciaceae with c. 300 species worldwide. A major missing link in understanding its global biogeography has been eastern Asia where the genus has never been systematically revised. Here we review specimen and literature records for Rinodina for north-eastern Asia (Russian Far East, Japan and the Korean Peninsula) and recognize 43 species. We describe two species, R. hypobadia and R. orientalis, as new to science. Rinodina hypobadia is distinguished by its pigmented hypothecium, Dirinaria-type ascospores and pannarin in both thallus and epihymenium. Rinodina orientalis is characterized by its erumpent apothecia that remain broadly attached, with discs sometimes becoming convex and excluding the thalline margins, ascospores belonging to the Physcia-type and secondary metabolites absent. Nine other species are reported from the region for the first time. These include R. dolichospora, R. freyi, R. metaboliza, R. sicula, R. subminuta and R. willeyi. Of particular biogeographical interest are three additional new records that have western North American–eastern Asian distributions: the corticolous species R. endospora, R. macrospora and R. megistospora. Six species have the better known eastern North American–eastern Asian distributions: R. ascociscana (syn. R. akagiensis, R. melancholica), R. buckii, R. chrysidiata, R. subminuta, R. tenuis (syn. R. adirondackii) and R. willeyi, and two have eastern North American–eastern Asian–European distributions: R. excrescens and R. moziana (syn. R. destituta, R. vezdae). Our study begins to close one of the largest gaps in our knowledge of circumboreal species distributions in Rinodina and, together with previous studies in North America and Europe, provides new insights into circumboreal crustose lichen biogeography. Rinodina cinereovirens (syn. R. turfacea var. cinereovirens) is also reported as new to North America.

The new species Xenolecia cataractarum Fryday is described from Campbell Island. It differs from X. spadicomma, the only other species of the genus, in having much smaller apothecia and ascospores, an olivaceously pigmented epihymenium (brown in X. spadicomma), and a thallus with a non-amyloid medulla and norstictic acid (amyloid medulla and confluentic acid in X. spadicomma). Xenolecia spadicomma is reported here from several localities on the Falkland Islands and three from the Región de Los Lagos, Chile, which are the first reports of this species since its description from Isla Wellington in the south-west of Patagonia in 1868. A full description of X. spadicomma is also provided.

Thin-layer chromatography (TLC) still enjoys widespread popularity among lichenologists as one of the fastest and simplest analytical strategies, today remaining the primary method of assessing the secondary product content of lichens. The pitfalls associated with this approach are well known as TLC leads to characterizing compounds by comparison with standards rather than properly identifying them, which might lead to erroneous assignments, accounting for the long-held interest in hyphenating TLC with dedicated identification tools. As such, commercially available TLC/Mass Spectrometry (MS) interfaces can be easily connected to any brand of mass spectrometer without adjustments. The spots of interest are extracted from the TLC plate to retrieve mass spectrometric signals within one minute, thereby ensuring accurate identification of the chromatographed substances. The results of this hyphenated strategy for lichens are presented here by 1) describing the TLC migration and direct MS analysis of single lichen metabolites of various structural classes, 2) highlighting it through the chemical profiling of crude acetone extracts of a set of lichens of known chemical composition, and finally 3) applying it to a lichen of unknown profile, Usnea trachycarpa.

In the present study, we characterized the mating-type locus of Knightiella splachnirima (Icmadophilaceae) using degenerate and inverse PCR techniques. We screened for the presence of both mating-type locus idiomorphs in DNA extracts of minuscule samples of haploid thalline tissue. We found that only one of the two idiomorphs was present in each sample, and the mating-type ratio (MAT1-1:MAT1-2) was very balanced, being 8:10 and 13:14 at local and global scales, respectively. This indicates that the species is probably self-incompatible and requires the presence of compatible mating partners for sexual reproduction (heterothallic mating system). Furthermore, we provide a mating-type screening protocol with K. splachnirima specific mating-type locus primers, which could serve as an essential tool for the conservation management of this rare Australasian endemic.

We assessed the taxonomy and phylogenetic relationships of Hawaiian species of the neotropical genus Lobariella. A single species was previously reported from the archipelago, the widespread neotropical L. crenulata. We targeted three loci of the mitochondrial and nuclear ribosomal rDNA gene cistron (mtSSU, nuLSU, ITS) and also performed a molecular clock analysis. Our results show that L. crenulata s. str. is seemingly present in the archipelago based on older herbarium collections. However, Hawaiian Lobariella includes three additional, presumably endemic, species unrelated to L. crenulata and new to science: L. flynniana Lücking, Moncada & C. W. Sm., with richly branched marginal phyllidia giving the thallus a fruticose appearance; L. robusta Lücking, Moncada & C. W. Sm., with a rather thick thallus and abundant, large, laminal phyllidia; and L. sandwicensis Lücking, Moncada & C. W. Sm., with a delicate thallus producing apothecia. Lobariella flynniana represents a novel morphotype within the genus, thus far known only from Hawaii. All three species are very closely related, forming a well-supported, monophyletic clade in spite of their morphological differences, suggesting local micro-radiation. Molecular clock analysis indicates that this clade colonized the islands between 1–8 mya and diverged between 0–2 mya. We interpret recent colonization as one of the main reasons why this clade has not diversified further.

Lichens as sessile and slow-growing symbiotic associations have evolved various carbon-based secondary compounds (CBSCs) to mitigate the effects of some stressors in the extreme environments in which they often grow. The mat-forming lichen Cladonia stellaris, an important fodder for reindeer, produces usnic acid in the outermost layer and perlatolic acid in the medulla. Here, we studied effects of simulated global warming on these CBSCs in C. stellaris cultivated in climate chambers with: 1) ambient conditions as control or 2) ambient conditions +4°C. The chambers simulated, at an hourly resolution, an averaged 10-year growing season dynamic from a long-term monitored boreal mire in northern Sweden. After two months of acclimation, +4°C warming in one simulated growing season increased the concentration of usnic acid by 31% compared with ambient conditions, whereas the warming decreased the concentration of perlatolic acid by 14%. Since lichen CBSCs play important roles in ecosystem processes such as lichenivory and decomposition, these changes may profoundly affect lichen-dominated ecosystems.