This Special Issue is dedicated to Professor Pier Luigi Nimis on the occasion of his 70th birthday and retirement. It was our aim to publish papers addressing the three major research fields that Pier Luigi dealt with during his career: systematics and taxonomy, biomonitoring and ecology, and data resources and digitization. The papers in this Special Issue provide a vivid overview of the state of the art in the three research fields, and they reflect on Pier Luigi's outstanding contribution to lichenology. They offer a wide array of different methodologies, from the traditional approaches investigating lichen diversity and taxonomy by means of morpho-anatomical analyses, culture isolations and phylogenetic systematics, to the most modern sequencing techniques, and to the development of computer-aided tools and databases for facilitating lichen identification.

There are people who leave their mark in their field. Without doubt, Pier Luigi Nimis (for the registry office, Pierluigi; for friends, Pier; for family members, Pil; for me and a few colleagues, PL) is such a person. On the threshold of retirement but no less active than ever, Pier Luigi is about to begin a new phase of his life, a life dedicated entirely to science and, in particular, to lichenology.

Urban green spaces are indispensable for the conservation of biodiversity in Germany. In addition, the availability of green areas often provides citizens with the only opportunity to experience nature. Lichens are an important component of urban environments in terms of both species diversity, and ecosystem functions and services. However, they are rarely the subject of biodiversity education. To bolster awareness on their diversity and appreciation, a transformative biodiversity education in both the formal and informal sectors is necessary. This transformative biodiversity education should not only provide knowledge about species and habitats, but also on all dimensions of biodiversity, viz., the three levels of biodiversity, drivers of biodiversity loss, and ecosystem services. For this reason, the design of biodiversity education may be particularly challenging for educators and teachers. This paper shows how biodiversity education projects on urban lichens can be developed in accordance with the principles of transformative education, supporting nature experience, knowledge transfer (species knowledge in a broad sense and interdisciplinary aspects), participation and cooperation, as well as the use of digital media. Two best-case projects, tested in Germany, are presented as examples for the design and implementation of a transformative lichen education in urban areas. A similar approach can be easily applied in other education systems beyond national boundaries.

Africa is an important global reservoir for biological, cultural and traditional knowledge about fungi and lichens, which are used as food, medicine and in mythology, among other things. African human populations are undergoing highly significant changes and adaptation processes, which are accompanied by rapid urbanization, meeting with western civilization, high rural migration and the loss of natural ecosystems. Indigenous knowledge is being lost, including that concerning fungi and lichens. Ethnomycology and ethnolichenology provide a diversity of knowledge about beneficial and poisonous fungi and lichens, and give insights into their sociological impact on human behaviour and use. Here we present a working and publishing environment established with the Diversity Workbench software in line with national and international initiatives for FAIR guided provision of research data. The database application called ‘EthnoMycAfrica’ contains published ethnomycological and ethnolichenological information from Africa. The content is created and curated by team partners from Central, East, West, North and Southern Africa. Data entry is performed both online and offline, optionally via a mobile device. Currently, the system with the tools DiversityDescriptions and DiversityNaviKey contains a total of 1350 well-structured and freely and openly accessible data records. EthnoMycAfrica is the first database with a data schema, standard descriptors and data content created mainly by African scholars. The data can be useful for researchers, students, conservationists, policy makers, and others. It will also provide a basis for facilitating hypothesis generation and meta-analysis.

We present a novel identification tool called PhyloKey, based on the method of morphology-based, phylogenetic binning developed within the software package RAxML. This method takes a reference data set of species for which both molecular and morphological data are available, computes a molecular reference tree, maps the morphological characters on the tree, and computes weights based on their level of consistency versus homoplasy using maximum likelihood (ML) and maximum parsimony (MP). Additional units for which only morphological data are known are then binned onto the reference tree, calculating bootstrap support values for alternative placements. This approach is modified here to work as an identification tool which uses the same character coding approach as interactive keys. However, rather than identifying individual samples through a progressive filtering process when entering or selecting characters, query samples are binned in batch mode to all possible alternative species in the tree, with each placement receiving a bootstrap support adding to 100% for all alternative placements. In addition to the fact that, after scoring a character matrix, a large number of specimens can be identified at once in short time, all possible alternative identifications are immediately apparent and can be evaluated based on their bootstrap support values. We illustrate this approach using the basidiolichen genus Cora, which was recently shown to contain hundreds of species. We also demonstrate how the PhyloKey approach can aid the restudying of herbarium samples, adding further value to these collections and contributing with large quantitative data matrices to ‘non-molecular museomics’. Our analysis showed that PhyloKey identifies species correctly with as low as 50% of the characters sampled, depending on the nature of the reference tree and the character weighting scheme. Overall, a molecular reference tree worked best, but a randomized reference tree gave more consistent results, whereas a morphological reference tree performed less well. Surprisingly, even character weighting gave the best results, followed by parsimony weighting and then maximum likelihood weighting.

iNaturalist is a widely-utilized platform for data collection and sharing among non-professional volunteers and is widely employed in citizen science. This platform's data are also used in scientific studies for a wide range of purposes, including tracking changes in species distribution, monitoring the spread of alien-invasive species, and assessing the impacts of urbanization and land-use change on biodiversity. Lichens, due to their year-round presence on trees, soil and rocks, and their diverse shapes and colours, have captured the attention of iNaturalist users, and lichen records are widely represented on the platform. However, due to the complexity of lichen identification, the use of data collected by untrained, or poorly trained volunteers in scientific investigation poses concerns among lichenologists. To address these concerns, this study assessed the reliability of lichen identification by iNaturalist users by comparing records on the platform with identifications carried out by experts (experienced lichenologists) in three cities where citizen science projects were developed. Results of this study caution against the use of unchecked data obtained from the platform in lichenology, demonstrating substantial inconsistency between results gathered by iNaturalist users and experts.

A checklist of Lichen-forming, Lichenicolous and Allied Fungi of Ecuador is presented with a total of 2599 species, of which 39 are reported for the first time from the country. The names of three species, Hypotrachyna montufariensis, H. subpartita and Sticta hypoglabra, previously not validly published, are validated. Pertusaria oahuensis, originally introduced by Magnusson as ‘ad interim’, is validated as Lepra oahuensis. The form Leucodermia leucomelos f. albociliata is validated. Two new combinations, Fissurina tectigera and F. timida, are made, and Physcia mobergii is introduced as a replacement name for the illegitimate P. lobulata Moberg non (Flörke) Arnold. In an initial step, the checklist was compiled by reviewing literature records of Ecuadorian lichen biota spanning from the late 19th century to the present day. Subsequently, records were added based on vouchers from 56 collections participating in the Consortium of Lichen Herbaria, a Symbiota-based biodiversity platform with particular focus on, but not exclusive to, North and South America. Symbiota provides sophisticated tools to manage biodiversity data, such as occurrence records, a taxonomic thesaurus, and checklists. The thesaurus keeps track of frequently changing names, distinguishing taxa currently accepted from ones considered synonyms. The software also provides tools to create and manage checklists, with an emphasis on selecting vouchers based on occurrence records that can be verified for identification accuracy. Advantages and limitations of creating checklists in Symbiota versus traditional ways of compiling these lists are discussed. Traditional checklists are well suited to document current knowledge as a ‘snapshot in time’. They are important baselines, frequently used by ecologists and conservation scientists as an established naming convention for citing species reported from a country. Compiling these lists, however, requires an immense effort, only to inadequately address the dynamic nature of scientific discovery. Traditional checklists are thus quickly out of date, particularly in groups with rapidly changing taxonomy, such as lichenized fungi. Especially in megadiverse countries, where new species and new occurrences continue to be discovered, traditional checklists are not easily updated; these lists necessarily fall short of efficiently managing immense data sets, and they rely primarily on secondary evidence (i.e. literature records rather than specimens). Ideally, best practices make use of dynamic database platforms such as Symbiota to assess occurrence records based both on literature citations and voucher specimens. Using modern data management tools comes with a learning curve. Systems like Symbiota are not necessarily intuitive and their functionality can still be improved, especially when handling literature records. However, online biodiversity data platforms have much potential in more efficiently managing and assessing large biodiversity data sets, particularly when investigating the lichen biota of megadiverse countries such as Ecuador.

Tremella caloplacae (Zahlbr.) Diederich is a species complex including at least nine different species. Here, we formally describe the new species Tremella elegantis, T. nimisiana, T. parietinae, T. pusillae and T. sorediatae. Tremella elegantis induces galls in the hymenium of Rusavskia elegans and forms 2-celled basidia, where cells rarely elongate and sometimes give the appearance of two immature, independent basidia. Tremella nimisiana has small basidiomata (less than 1 mm diam.), narrowly ellipsoid to pyriform 2-celled, occasionally clavate to subcylindrical 3-celled basidia, and grows in the hymenium of Xanthocarpia species. Tremella parietinae is characterized by the exclusive growth in the hymenium of Xanthoria parietina, the broadly fusiform to ellipsoid probasidia, and the subspherical, pyriform or ellipsoid 2(–3)-celled basidia. Tremella pusillae has ellipsoidal probasidia, 2(–3)-celled pyriform or ellipsoidal basidia that sometimes are constricted at the septum, and grows only on Calogaya pusilla. Tremella sorediatae is characterized by inducing galls on the thallus of Rusavskia sorediata and by pyriform to ellipsoid basidia that sometimes are constricted at the septum. Three species are not formally described and are left unnamed as Tremella sp. 13 on Calogaya biatorina, Tremella sp. 14 on Calogaya decipiens and Tremella sp. 15 on Polycauliona sp. Tremella caloplacae in the strict sense is re-circumscribed as a species confined to Variospora species.

Arthonia epipolytropa Hafellner & Grube and Arthonia subclemens Hafellner, Grube & Muggia are described as new to science. Both are specific parasites of Lecanora polytropa s. lat., but of differing pathogenicity and of very different appearance. Whereas the clearly parasitic Arthonia epipolytropa with its agglomerated ascomata is presently known with certainty from a number of localities along the arch of the Alps (Austria, Italy, Switzerland), other parts of Europe (Norway, Albania) and northern America (USA), the less harmful A. subclemens with its isolated sunken ascomata is currently known only from a small number of localities in the Eastern Alps (Austria, Italy) and various mountain ranges in southern Europe (Spain, North Macedonia, Greece). The species are compared with other Arthonia species known from Lecanora or one of its recently segregated genera. A key to the fungi regularly found on Lecanora polytropa s. lat. is presented.

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.