Species of lichen-forming fungi (LFF) display an array of geographical distribution patterns. Among the broadly distributed lichen-forming fungal species, the degree of reproductive isolation and genetic substructure among populations varies widely, in some cases masking unrecognized diversity or meaningful biogeographical patterns. Lecidea atrobrunnea (Raymond ex Lam. & DC.) Schaer. s. lat. (Lecideaceae) is a widespread species complex that has been studied for over two centuries since its initial description. The diversity of the L. atrobrunnea group is highest in western North America, where a dizzying array of morphologies and chemistry can occur at local scales. Here we investigate whether the assumed cosmopolitan distribution of L. atrobrunnea s. lat. is an artifact of taxonomic limitations and masks biogeographical patterns in this species complex. To address these questions, we compiled sequence data from the standard fungal barcoding marker (ITS) for over 100 specimens within this complex, in addition to genome-scale data from a subset of these representing over 1600 single-copy nuclear genes spanning over 3 Mb of the genome. Our study corroborates the perspective that the morphologically and chemically variable Lecidea atrobrunnea group reflects a complex of distinct species-level lineages, with 42–83 candidate species inferred from the ITS region and high levels of diversity inferred from a subset of specimens using genome-scale data. However, both phenotype- and molecular-based species boundaries remained unsettled, with the most common nominal taxa recovered as highly polyphyletic and with conflict among different molecular species delimitation approaches. Our study also highlights the potential for geographically restricted species, with fascinating biogeographical patterns, challenging, in part, the assumed cosmopolitan distribution of L. atrobrunnea s. lat. This study provides valuable direction for future research that will be crucial in understanding diversification and establishing a robust taxonomy for this well-known species complex.

There is still a high diversity of lichen-forming fungi that remains undescribed, especially cryptic lineages at the species level. Integrating morphological, chemical, and DNA sequence data has proved useful in corroborating species descriptions and delimitations. Here we reviewed morphological features, secondary metabolites and the DNA sequences of ITS, mtSSU and nuLSU markers to study the diversity of Xanthoparmelia in southern Africa. A total of 37 species were recorded. Three of these appear undescribed, and we therefore describe them here as new: Xanthoparmelia nimisii, with a sorediate thallus and broad lobes, is well supported as a clade separate from X. annexa; X. pseudochalybaeizans with a white medulla is phylogenetically distinct from the otherwise similar X. chalybaeizans; and X. sipmaniana, well supported as a separate clade from the similar X. hypoprocetrarica. In addition, the separation of Xanthoparmelia capensis and X. tinctina requires further studies.

During a study of the incompletely known lichen flora of the Caucasus, we analyzed 237 specimens of corticolous Bacidia s. str. collected in the Northern and Southern Caucasus, including Armenia, Azerbaijan, Georgia, and Russia. Of these, 54 specimens belonging to 11 species of Bacidia s. str. were selected for molecular studies, representing the observed morphological variability of the genus. We obtained 142 sequences from three RNA-coding genes (nrITS, nrLSU, and mtSSU) and two protein-coding genes (RPB1 and RPB2). The single and concatenated datasets were complemented with Bacidia s. str. sequences from GenBank and subjected to Bayesian inference and two maximum likelihood analyses (RAxML and IQ-TREE). The resulting trees yielded highly concordant topologies of the groups and corresponded with previous results, supporting two main clades correlating with apothecia pigmentation. Our analyses are the first to reveal the presence of Bacidia heterochroa in the Caucasus. An exceptionally high degree of morphological plasticity was found in the Rubella and Suffusa groups. As a result of morphological examination and phylogenetic results, B. caucasica (Suffusa group) was described as new to science. Furthermore, two putative taxa in the Rubella group, Bacidia inconspicua ined. and B. maritima ined., were introduced. This study furthers our understanding and documentation of the understudied lichen flora of the Caucasus, bringing the total number of Bacidia species for the region to 13.

Five species of Schaereria Körb. are known to occur in Australia (including Tasmania). The new species S. australis Kantvilas is described from alpine Tasmania and New South Wales; it is characterized by a thallus lacking lichen substances, apothecia with only brown pigments, and ellipsoid, uniseriate ascospores, 10−17 × 6−9 μm. Also treated are: S. bullata Kantvilas, endemic to Tasmania; S. dolodes (Nyl. ex Hasse) Schmull & T. Sprib., first described from North America and recorded here for the first time from the Southern Hemisphere (Tasmania); the bipolar S. fuscocinerea (Nyl.) Clauzade & Cl. Roux and the Australian endemic S. xerophila Rambold & H. Mayrhofer, both recorded for the first time from Tasmania. The species are illustrated and an identification key is provided.

Our floristic work in British ancient forests resulted in a description of a frequently reported but misidentified species, Coenogonium nimisii. Its thallus is very similar to Porina rosei, but the apothecia and pycnidia correspond with C. luteum. Sterile collections are not easy to distinguish but the new species differs from P. rosei in several microscopic characters of the isidia. Coenogonium nimisii is so far known from bark and epiphytic bryophytes, rarely mossy rocks, in ancient humid forests of Great Britain and Ireland. The genus Coenogonium is poorly represented by molecular data in the GenBank database. Our preliminary results revealed distinct genetic lineages within two traditionally circumscribed species, C. luteum and C. pineti, which may represent cryptic species.

Peltigera globulata Miadl. & Magain, a new species in the P. ponojensis/monticola species complex of section Peltigera, is formally described. This clade was previously given the interim designation Peltigera sp. 17. It is found in sun-exposed and xeric habitats at high altitudes in Peru and Ecuador. Peltigera globulata can be easily recognized by its irregularly globulated margins covered mostly by thick, white pruina, somewhat resembling the sorediate thallus margins of P. soredians, another South American species from section Peltigera. The hypervariable region of ITS1 (ITS1-HR), which is in general highly variable among species of section Peltigera, does not have diagnostic value for species identification within the P. ponojensis/monticola complex. Nevertheless, no significant level of gene flow was detected among eight lineages representing a clade of putative species (including P. globulata) within this complex. ITS sequences from the holotype specimens of P. monticola Vitik. (collected in 1979) and P. soredians Vitik. (collected in 1981) and lectotype specimens of P. antarctica C. W. Dodge (collected in 1941) and P. aubertii C. W. Dodge (collected in 1952) were successfully obtained through Sanger and Illumina metagenomic sequencing. BLAST results of these sequences revealed that the type specimen of P. monticola falls within the P. monticola/ponojensis 7 clade, which represents P. monticola s. str., and confirmed that the type specimen of P. aubertii falls within a clade identified previously as P. aubertii based on morphology. The ITS sequence from the type specimen of P. soredians, which superficially resembles P. globulata, confirms its placement in the P. rufescens clade. Finally, we discovered that the name P. antarctica was erroneously applied to a lineage in the P. ponojensis/monticola clade. The ITS sequence from the type specimen of P. antarctica represents a lineage within the P. rufescens clade, which is sister to the P. ponojensis/monticola clade.

A unique crustose lichen species was recently documented from various types of preserved forests across boreal and temperate Europe (Norway, Ukraine, the Czech Republic) and the Caucasus (Russia). It is formally described here as the new species Biatora amylacea. A phylogeny based on ITS and mtSSU sequences demonstrates that it belongs to an isolated group within the core of Biatora s. lat., together with the recently described B. radicicola. It is a distinctive taxon within the genus on account of its amyloid exciple, otherwise known only from members of the Biatora rufidula group. The new species is also characterized by amyloid thalline hyphae and the production of soredia with a blue-green pigment. This microlichen may serve as a bioindicator species of old-growth forests.

The new genus Nimisora Pérez-Ort., M. Svenss. & J. C. Zamora is introduced to accommodate a puzzling lecideoid epiphyte common in the central Iberian Peninsula. Nimisora is characterized by the following combination of characters: lecideoid apothecia, excipulum composed of sparingly branched radiating hyphae with narrow lumina, thick walls and swollen terminal cells, the presence of a brown K+ olivaceous green pigment in the epihymenium, an ascus tip similar to the Bacidia-type, and the presence of simple ellipsoid ascospores. Molecular analyses based on nrITS, nrLSU and mtSSU sequences unequivocally place the new genus within the Lecanoraceae; however, its phylogenetic affinities with other genera of the family remain largely unresolved. Comparisons with the morphologically closest genera are provided. The single species of the genus, Nimisora iberica Pérez-Ort., Turégano, M. Svenss. & J. C. Zamora sp. nov., is also described as new to science.

Porina is a widely distributed, species-rich genus of crustose, lichen-forming fungi, some with thalline outgrowths that have been recognized as isidia. We studied three taxa with thalli consisting chiefly of ascending isidioid structures occurring on trunks and branches of Taxodium in southwestern Florida, and provide details of their structure with light and electron microscopy. Two of these taxa we describe as new species: P. microcoralloides and P. nanoarbuscula. Genetic sequences (mtSSU) suggest that they are closely related to each other, yet they differ markedly in the size, morphology and anatomical organization of their isidioid branches as well as in the length of their ascospores. In the three Floridian taxa studied, the crustose portion of the thallus is partly endophloeodic and partly superficial, the latter often patchy, evanescent or inconspicuous, and completely lacks the differentiated anatomical organization characteristic of the isidioid structures arising from it. In Porina microcoralloides, the ascendant thallus consists of branched, coralloid inflated structures with phycobiont (Trentepohlia) unicells arranged at the periphery of a loose central medulla. Sparse fungal cells are interspersed and overlie the algal layer in places, but no differentiated cortex is present, leaving phycobiont cells more or less exposed at the surface. In the closely related Porina nanoarbuscula, the isidioid structures are much finer, more densely branched, and composed of a single, central file of roughly spherical Trentepohlia cells surrounded by a jacket of subglobose fungal cells. The ascospores of P. microcoralloides are more than twice the length of those of P. nanoarbuscula. Although thalli of these two Porina species occur in the same habitats and are sometimes found growing alongside each other, phylogenetic analysis of rbcL sequences suggest that they partner with distinct clades of Trentepohlia phycobionts. A third taxon examined, Porina cf. scabrida, is morphologically rather similar to P. microcoralloides, but the ascendant branches are bright yellow-orange, more cylindrical, and corticated by a thin layer of agglutinated fungal hyphae; perithecia were not seen. Analysis of mtSSU sequences places it distant from P. microcoralloides and P. nanoarbuscula phylogenetically. None of the Floridian taxa studied was particularly close to the European isidiate species Porina hibernica and P. pseudohibernica, which appeared as sister to each other in the analysis. While a particular type of isidiose structure may be reliably characteristic of specific taxa, similarities or differences in these structures do not seem to be useful indicators of phylogenetic proximity or distances among taxa. The morphological trends evident in Porina suggest that multiple transitions from crustose to isidioid or microfruticose growth have arisen repeatedly and in quite different ways within this single genus. At least some of the diverse structures treated within the broad concept of isidia may be representative of the developmental pathways by which fruticose growth forms may arise.

The manna lichens, a group of vagrant species with subfruticose and subfoliose thalli in the genus Circinaria Link, have received attention for millennia. Here, a new manna lichen species, Circinaria nimisii sp. nov. (Megasporaceae), is described and illustrated. This vagrant lichen is found on Mount Olympus in Greece and is the fourth known manna lichen in Europe. The new taxon is characterized by its subfruticose, densely-branched thallus with a muddy, earthy colour, whitish pseudocyphellae on tips of branches, mature apothecia distinctly adnate to stipitate, and paraplectenchymatous cortex tissue. Molecular sequence data from the standard barcoding marker (nrITS) also corroborate the distinction of this species from closely related congeners. Finally, Agrestia zerovii, previously known only from its type locality in Ukraine, is proposed as a new synonym of Circinaria hispida.

Three twig-growing lichen species belonging to the family Teloschistaceae from southern Patagonia are described as new to science: Marchantiana pyramus sp. nov., with minute apothecia, orange due to dominant content of emodin, M. ramulicola sp. nov., with minute olive apothecia with dominant parietin and Austroplaca thisbe sp. nov., with clear yellow apothecia also with dominant parietin. Marchantiana subpyracea, M. epibrya and M. queenslandica are new combinations for species which, so far, are known only from New Zealand and Australia. Blastenia circumpolaris is shown to be very common in Patagonia and Marchantiana asserigena is documented for the first time from the Southern Hemisphere, viz. the Falkland Islands. The genus Marchantiana is analyzed here using three genes and is shown to be closely related to Yoshimuria; although appearing as paraphyletic, a monophyletic origin cannot be ruled out. Morphology, ecology and distribution support a monophyletic treatment and Marchantiana is therefore treated as such.

An increasing number of studies are describing the diversity of lichen phycobionts, which is leading to a better understanding of how lichen communities are assembled at different taxonomic, evolutionary and geographical scales. The present study explores the identity and genetic diversity of the microalgal partners of Punctelia borreri and P. subrudecta, two tropical and temperate parmelioid lichen fungi that often grow in temperate and Mediterranean forest ecosystems in Europe. Based on a specimen sampling distributed in two climatically divergent regions in the Iberian Peninsula, we found that these mycobionts are associated with Trebouxia gelatinosa, whose identity was also confirmed by an ultrastructural study of the pyrenoid. The bipartite network analysis indicated that each Punctelia species was associated with a different set of low frequency T. gelatinosa infraspecific lineages, whereas the two most abundant phycobiont lineages were shared between both mycobionts. Based on the current sampling, these two algal lineages occur exclusively in one of the two studied regions, which might point towards climate-driven, fine-tuned fungal-algal interactions. Finally, we documented visible symptoms of injury on the thalli in areas likely to have been impacted by air pollution.

Species can respond to climate change by migrating to track their suitable climate space, and/or through adaptation (across generations) or acclimation (by individuals) to a changed in situ environment. Lichens provide an excellent model for studying acclimation; being poikilohydric, there is strong evidence that their phenotype presents an adaptation to different moisture regimes, and that key aspects of the phenotype, notably specific thallus mass (STM), have plasticity towards effective acclimation that maximizes water storage in drier environments. In this study we quantified acclimation of STM for Lobaria pulmonaria across a regional climatic gradient, and within sites for different microclimates, using a one-year common garden growth experiment. We found that STM tended to increase with thallus growth; however, when accounting for growth, STM shifted to be lower than average in wetter environments, higher than average in intermediate environments, and failed to respond in the driest environment where growth was compromised. The possibility of phenotypic acclimation in Lobaria pulmonaria appears to be functionally linked to the propensity for growth, and we present a scheme coupling growth with STM to define the limits of the species realized niche.

Environmental control strategies are commonly practised to limit biodeterioration issues threatening indoor cultural heritage objects, while they are still poorly exploited for the conservation of outdoor stone heritage surfaces, including rock art. In this study, we evaluated the environmental factors driving the diversity and abundance of lithobiontic communities in the Rock Engravings National Park of Naquane (UNESCO WHS n. 94, Italy). The survey considered 23 rocks that had been cleaned in the last three (3YC) or twelve (12YC) years or more than 40 years ago (NRC). A cyanobacteria-dominated biofilm and lichens (37 taxa) were the most widespread and abundant lithobiontic components, prevailing on 3YC–12YC and NRC rocks, respectively. On the latter, a turnover of xerophytic and meso-hygrophytic lichen communities was observed. On 3YC–12YC rocks lichen colonization, if present, was limited to nitrophytic species, including common epiphytes from surrounding trees and a small number of meso-hygrophytic species, with a prevalence of asexual reproductive strategies. Multivariate analyses including environmental parameters (canonical correspondence analyses) indicated that tree cover and the presence of bare or vegetated ground upstream of the rocks, probably prolonging wetness and providing nutrients by water transport, are the factors mostly related to the microbial and lichen recolonization of 3YC–12YC surfaces. On this basis, an experiment on preventive conservation was conducted, consisting of a new cleaning of a strongly recolonized 3YC surface combined with the building of a small wall to protect part of the rock from prolonged water fluxes. The fluorimetric and colorimetric monitoring of the rock surface, carried out 40 months after this new cleaning intervention, displayed recolonization on the unprotected area only, indicating the potential of preventive conservation strategies in outdoor environments.

Although epiphytic lichens are widely adopted as environmental indicators, they are not yet included among the target species listed in Annex II of the Habitats Directive, to which the system of protected areas of the Natura 2000 network refers. In this work, we aim to test the effectiveness of this system, mainly designed for the conservation of other groups of species, in protecting lichen species richness. For this purpose, we considered a case study (Central Italy) with half of the territory included in protected areas. Statistical differences in species richness and lichen communities were tested between sites located in 16 Protected Areas (PA) and 11 Non-Protected Areas (NPA) using non-parametric tests, multi-response permutation procedures (MRPP), non-metric multidimensional scaling (NMDS) and Indicator Species Analysis (ISA). Despite the broad overlap between epiphytic lichen communities of NPAs and PAs and a similar number of total and common species, PAs contain a significantly higher number of nationally rare and extremely rare species, including cyanolichens. These results are also confirmed by the indicator analysis. Although the Natura 2000 network does not explicitly address the conservation of lichens, the protected areas in our study can play a role in protecting the diversity of epiphytic lichens, especially nationally rare and endangered species. However, the future inclusion of red-listed epiphytic lichens among the target species of Annex II of the Habitats Directive would be welcome to better protect these organisms on a European level.

The species–area relationship (SAR) states that species richness increases with the increase of the sampled area, although other factors can influence the pattern. SARs have been tested on many different organisms, but only rarely on lichens. We aimed to test the SAR, across a wide range of area sizes, for three main substratum-related guilds of lichens, namely epiphytic, epilithic and epigaeic. The test was performed using data from lichen inventories carried out in 44 protected areas of various sizes across Italy. We found a positive correlation of species richness with area size for all three guilds, better fitted by the logarithmic function for epilithic lichens and by the power function for epiphytic and epigaeic lichens. Our results support the fundamental role of area size as the main driver for lichen diversity, suggesting that in an area-based conservation framework, larger protected areas are fundamental to support high lichen species richness. However, finer scale investigations are also required to better elucidate whether and how other environmental factors could interact with area size and modify SAR patterns. Exhaustive lichen inventories could be useful information sources to more robustly test such relationships, and therefore better inform conservation practices.