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Resumen Presentamos una actualización nomenclatural y taxonómica del Catálogo de Liqúenes de Colombia. Como resultado, el número de nombres reportados se reduce de 1821 a 1793, el total de taxones aceptados de 1732 a 1675, y el número de especies de 1672 a 1634. El número de reportes dudosos o excluidos, que incluye nombres con estado nomenclatural o taxonómico no resuelto, se aumenta de 80 a 109. El número de géneros reportados para Colombia aumenta de 272 a 306; se excluyen 29 géneros anteriormente reportados y se incluyen 63 géneros hasta la fecha no reportados. La actualización reduce el total de familias de 73 a 69; resulta en la eliminación de 13 familias y la inclusión de 9 familias anteriormente no reportadas. Basado en la revisión de material antes reportado bajo nombres incorrectos, se reportan tres nuevos registros para Colombia: Caloplaca granularis (Müll.Arg.) Zahlbr., Haematomma persoonii (Fée) A. Massal. y Ochrolechia subpallescens Verseghy. Además, confirmamos la presencia de las especies: Physcia crispula Müll.Arg., Pseudocyphellaria citrina (Gyeln.) Lücking, Moncada & S. Stenroos, Pseudocyphellaria sandwicensis (Zahlbr.) Moncada & Lücking, Pseudocyphellaria xanthosticta (Pers.) Moncada & Lücking, Sticta sylvatica (Huds.) Ach., Usnea crenulata Truong & P. Clerc y Usnea mexicana Vain. También se introducen 16 novedades nomenclaturales: Ancistrosporella gracilior (Nyl.) Lücking comb. nov. (basiónimo: Opegrapha gracilior Nyl.); Bacidia neofusconigrescens Lücking nom. nov. (sinónimo reemplazado: Lecidea millegrana var. fusconigrescensNyl.) [non Bacidiafusconigrescens (Kremp.) Zahlbr.]; Diploschistes bartlettii (Lumbsch) Lücking comb. et stat. nov. (basiónimo: Diploschistes muscorum subsp. bartlettii Lumbsch); Gymnographopsis koreaiensis (Sipman) Lücking & Sipman comb. nov. (basiónimo: Graphis koreaiensis Sipman); Imshaugia angustior (Nyl.) Sipman (basiónimo: Parmelia angustior Nyl.); Kalbographa cabbalistica (Nyl.) Lücking comb. nov. (basiónimo: Graphis cabbalistica Nyl.; sinónimo nuevo: Graphina caracasana Müll.Arg.); Leptogium pseudolivaceum Lücking nom. nov. (sinónimo reemplazado: Collema olivaceum Hook.) [nom. illeg., non Leptogium olivaceum F. Wilson]; Malmidea demutans (Nyl.) Lücking comb. nov. (basiónimo: Lecidea demutans Nyl.); Ocellularia leucocarpoides (Nyl.) Lücking comb. nov. (basiónimo: Thelotrema leucocarpoides Nyl.; sinónimo nuevo: Ocellularia fuscospora Lücking & Pérez-Ort.); Phaeographis decolorascens (Nyl.) Lücking comb. nov. (basiónimo: Graphis decolorascens Nyl.); Phlyctis endecamera (Nyl.) Lücking & Sipman comb. nov. (basiónimo: Platygrapha endecamera Nyl.); Sprucidea fuscula (Nyl.) Lücking comb. nov. (basiónimo: Lecideafuscula Nyl.); Sticta rudiuscula (Vain.) Moncada & Lücking comb. et stat. nov. (basiónimo: Sticta damicornis f. rudiuscula Vain.); Sticta subdenudata Moncada & Lücking nom. nov. (sinónimo reemplazado: Sticta laciniata var. denudata Nyl.) [non Sticta denudata Taylor]; Thalloloma scribillans (Nyl.) Lücking comb. nov. (basiónimo: Graphis scribillans Nyl.; sinónimo nuevo: Graphis anguiniformis Vain.); y Yoshimuriella denudata (Taylor) Moncada & Lücking comb. nov. (basiónimo: Sticta denudata Taylor). Abstract We present a nomenclatural and taxonomic update of the Catálogo de Liquenes de Colombia. As a result, the number of reported names is reduced from 1821 to 1793, the total of accepted taxa from 1732 to 1675, and the number of species from 1672 to 1634. The number of doubtful or excluded reports, which includes names with unresolved nomenclatural or taxonomic status, increases from 80 to 109. The number of genera reported for Colombia increases from 272 to 306; 29 previously reported genera are excluded, and 63 genera not reported previously are included. The update reduces the total of families from 73 to 69; it results in the elimination of 13 families and the addition of 9 previously unreported families. We include three new records for Colombia: Caloplaca granularis (Müll.Arg.) Zahlbr., Haematomma persoonii (Fée) A. Massal., and Ochrolechia subpallescens Verseghy. Furthermore, we confirm the presence of the following species: Physcia crispula Müll. Arg., Pseudocyphellaria citrina (Gyeln.) Lücking, Moncada & S. Stenroos, Pseudocyphellaria sandwicensis (Zahlbr.) Moncada & Lücking, Pseudocyphellaria xanthosticta (Pers.) Moncada & Lücking, Sticta sylvatica (Huds.) Ach., Usnea crenulata Truong & P. Clerc, and Usnea mexicana Vain. We also introduce 16 nomenclatural novelties: Ancistrosporella gracilior (Nyl.) Lücking comb. nov. (bas.: Opegrapha gracilior Nyl.); Bacidia neofusconigrescens Lücking nom. nov. (replaced syn.: Lecidea millegrana var. fusconigrescens Nyl.) [non Bacidia fusconigrescens (Kremp.) Zahlbr.]; Diploschistes bartlettii (Lumbsch) Lücking comb. et stat. nov. (bas.: Diploschistes muscorum subsp. bartlettii Lumbsch); Gymnographopsis koreaiensis (Sipman) Lücking & Sipman comb. nov. (bas.: Graphis koreaiensis Sipman); Imshaugia angustior (Nyl.) Sipman (bas.: Parmelia angustior Nyl.); Kalbographa cabbalistica (Nyl.) Lücking comb. nov. (bas.: Graphis cabbalistica Nyl.; new syn.: Graphina caracasana Müll.Arg.); Leptogium pseudolivaceum Lücking nom. nov. (replaced syn.: Collema olivaceum Hook.) [nom. illeg., non Leptogium olivaceum F. Wilson]; Malmidea demutans (Nyl.) Lücking comb. nov. (bas.: Lecidea demutans Nyl.); Ocellularia leucocarpoides (Nyl.) Lücking comb. nov. (bas.: Thelotrema leucocarpoides Nyl.; new syn.: Ocellularia fuscospora Lücking & Pérez-Ort.); Phaeographis decolorascens (Nyl.) Lücking comb. nov. (bas.: Graphis decolorascens Nyl.); Phlyctis endecamera (Nyl.) Lücking & Sipman comb. nov. (bas.: Platygrapha endecamera Nyl.); Sprucidea fuscula (Nyl.) Lücking comb. nov. (bas.: Lecidea fuscula Nyl.); Sticta rudiuscula (Vain.) Moncada & Lücking comb. et stat. nov. (bas.: Sticta damicornis f. rudiuscula Vain.); Sticta subdenudata Moncada & Lücking nom. nov. (replaced syn.: Sticta laciniata var. denudata Nyl.) [non Sticta denudata Taylor]; Thalloloma scribillans (Nyl.) Lücking comb. nov. (bas.: Graphis scribillans Nyl.; new syn.: Graphis anguiniformis Vain.); and Yoshimuriella denudata (Taylor) Moncada & Lücking comb. nov. (bas.: Sticta denudata Taylor).

Lichens are symbiotic associations resulting from interactions among fungi (primary and secondary mycobionts), algae and/or cyanobacteria (primary and secondary photobionts), and specific elements of the bacterial microbiome associated with the lichen thallus. The question of what is a species, both concerning the lichen as a whole and its main fungal component, the primary mycobiont, has faced many challenges throughout history and has reached new dimensions with the advent of molecular phylogenetics and phylogenomics. In this paper, we briefly revise the definition of lichens and the scientific and vernacular naming conventions, concluding that the scientific, Latinized name usually associated with lichens invariably refers to the primary mycobiont, whereas the vernacular name encompasses the entire lichen. Although the same lichen mycobiont may produce different phenotypes when associating with different photobionts or growing in axenic culture, this discrete variation does not warrant the application of different scientific names, but must follow the principle "one fungus = one name". Instead, broadly agreed informal designations should be used for such discrete morphologies, such as chloromorph and cyanomorph for lichens formed by the same mycobiont but with either green algae or cyanobacteria. The taxonomic recognition of species in lichen-forming fungi is not different from other fungi and conceptual and nomenclatural approaches follow the same principles. We identify a number of current challenges and provide recommendations to address these. Species delimitation in lichen-forming fungi should not be tailored to particular species concepts but instead be derived from empirical evidence, applying one or several of the following principles in what we call the LPR approach: lineage (L) coherence vs. divergence (phylogenetic component), phenotype (P) coherence vs. divergence (morphological component), and/or reproductive (R) compatibility vs. isolation (biological component). Species hypotheses can be established based on either L or P, then using either P or L (plus R) to corroborate them. The reliability of species hypotheses depends not only on the nature and number of characters but also on the context: the closer the relationship and/or similarity between species, the higher the number of characters and/or specimens that should be analyzed to provide reliable delimitations. Alpha taxonomy should follow scientific evidence and an evolutionary framework but should also offer alternative practical solutions, as long as these are scientifically defendable. Taxa that are delimited phylogenetically but not readily identifiable in the field, or are genuinely cryptic, should not be rejected due to the inaccessibility of proper tools. Instead, they can be provisionally treated as undifferentiated complexes for purposes that do not require precise determinations. The application of infraspecific (gamma) taxonomy should be restricted to cases where there is a biological rationale, i.e., lineages of a species complex that show limited phylogenetic divergence but no evidence of reproductive isolation. Gamma taxonomy should not be used to denote discrete phenotypical variation or ecotypes not warranting the distinction at species level. We revise the species pair concept in lichen-forming fungi, which recognizes sexually and asexually reproducing morphs with the same underlying phenotype as different species. We conclude that in most cases this concept does not hold, but the actual situation is complex and not necessarily correlated with reproductive strategy. In cases where no molecular data are available or where single or multi-marker approaches do not provide resolution, we recommend maintaining species pairs until molecular or phylogenomic data are available. This recommendation is based on the example of the species pair Usnea aurantiacoatra vs. U. antarctica, which can only be resolved with phylogenomic approaches, such as microsatellites or RADseq. Overall, we consider that species delimitation in lichen-forming fungi has advanced dramatically over the past three decades, resulting in a solid framework, but that empirical evidence is still missing for many taxa. Therefore, while phylogenomic approaches focusing on particular examples will be increasingly employed to resolve difficult species complexes, broad screening using single barcoding markers will aid in placing as many taxa as possible into a molecular matrix. We provide a practical protocol how to assess and formally treat taxonomic novelties. While this paper focuses on lichen fungi, many of the aspects discussed herein apply generally to fungal taxonomy. The new combination Arthonia minor (Lücking) Lücking comb. et stat. nov. (Bas.: Arthonia cyanea f. minor Lücking) is proposed.

Aim: Phylogenetic diversification is a precursor to speciation, but the underlying patterns and processes are not well-studied in lichens. Here we investigate what factors drive diversification in two tropical, morphologically similar macrolichens that occupy a similar range but differ in altitudinal and habitat preferences, testing for isolation by distance (IBD), environment (IBE), and fragmentation (IBF). Location: Neotropics, Hawaii, Macaronesia. Taxon: Sticta andina, S. scabrosa (Peltigeraceae). Methods: We analysed 395 specimens from 135 localities, using the fungal ITS barcoding marker to assess phylogenetic diversification, through maximum likelihood tree reconstruction, TCS haplotype networks, and Tajima's D. Mantel tests were employed to detect structure in genetic vs. geographic, environmental, and fragmentation distances. Habitat preferences were quantitatively assessed by statistical analysis of locality-based BIOclim variables. Results: Sticta andina exhibited high phenotypic variation and reticulate phylogenetic diversity across its range, whereas the phenotypically uniform S. scabrosa contained two main haplotypes, one unique to Hawaii. Sticta andina is restricted to well-preserved andine forests and paramos, naturally fragmented habitats due to disruptive topology, whereas S. scabrosa thrives in lowland to lower montane zones in exposed or disturbed microsites, representing a continuous habitat. Sticta scabrosa showed IBD only across its full range (separating the Hawaiian population) but not within continental Central and South America, there exhibiting a negative Tajima's D. Sticta andina did not exhibit IBD but IBE at continental level and IBF in the northern Andes. Main conclusions: Autecology, particularly preference for either low or high altitudes, indirectly drives phylogenetic diversification. Low diversification in the low altitude species, S. scabrosa, can be attributed to rapid expansion and effective gene flow across a more or less continuous niche due to disturbance tolerance. In contract, high diversification in the high altitude species, S. andina, can be explained by niche differentiation (IBE) and fragmentation (IBF) caused by the Andean uplift.

El objetivo de este trabajo fue documentar la diversidad de líquenes cortícolas en el Chocó biogeográfico del Valle del Cauca, con descripciones breves de especies encontradas a lo largo de un gradiente altitudinal (0-3650 m). Se recolectaron ca. 3000 ejemplares, pertenecientes a 39 familias, 126 géneros y 750 especies. Las formas dominantes fueron las costrosas (1963 ejemplares), seguidas por las foliosas (719 ejemplares), las fruticosas (328 ejemplares) y las escuamulosas (168 ejemplares), mientras que las menos frecuentes fueron las gelatinosas y las pulverulentas. Las familias con mayor número de especies fueron Graphidaceae (188 especies), Parmeliaceae (97 especies), Arthoniaceae (53 especies), Lobariaceae (43 especies), Pyrenulaceae (40 especies), Ramalinaceae (38 especies) y Trypetheliaceae (30 especies), mientras que ocho familias solo presentaron una especie. Se reportan por primera vez cuatro géneros para Colombia: Clypeopyrenis, Distopyrenis, Megalotremis y Myeloconis.

Sticta is a subcosmopolitan genus most diverse in the tropics. Traditionally, many taxa were considered to be widespread and morphologically variable, following broadly circumscribed morphodemes. Among these is the S. weigelii morphodeme, characterized by a cyanobacterial photobiont and rather narrow, flabellate to truncate or tapering lobes producing predominantly marginal isidia. Molecular phylogenetic analyses focusing on the ITS fungal barcoding marker revealed that this morphodeme represents several species, some of which are only distantly related to each other. Here we describe two species and one subspecies of this morphodeme as new to science, based on analysis of 400 specimens, for 344 of which we generated ITS barcoding data. The two new species, S. andina and S. scabrosa, are broadly distributed in the Neotropics and also found in Hawaii, where the latter is represented by the new subspecies, S. scabrosa subsp. hawaiiensis; in the case of S. andina, the species is also found in the Azores. Sticta andina exhibits high phenotypic variation and reticulate genetic diversification, whereas the phenotypically rather uniform S. scabrosa contains two main haplotypes, one restricted to Hawaii. Sticta andina occurs in well-preserved montane to andine forests and paramos, whereas the two subspecies of S. scabrosa are found in tropical lowland to lower montane forests, tolerating disturbance and extending into anthropogenic habitats.Citation: Moncada B., Mercado-Diaz J. A., Smith C. W., Bungartz F., Serusiaux E., Lumbsch H. T. & Lucking R. 2021: Two new common, previously unrecognized species in the Sticta weigelii morphodeme (Ascomycota: Peltigeraceae). – Willdenowia 51: 35–45.Version of record first published online on 24 February 2021 ahead of inclusion in April 2021 issue.

We analyzed the structure of foliicolous lichen communities in the northernmost lowland forest of the Neotropics, Los Tuxtlas Tropical Biology Station in Veracruz, Mexico, and its dependence on phorophyte and microclimate. Along a 420‐m long transect with 15 equidistant sampling points, within a 10 m radius of each point, we sampled a total 137 phorophytes and 411 leaves. The phorophytes represented 13 species, with diverse leaf traits regarding size, texture, presence of hairs and/or glands, and longevity, including: Astrocaryum mexicanum (Arecaceae), Chamaedorea ernesti‐augustii (Arecaceae), Costus scaber (Costaceae), Guarea glabra (Meliaceae), Heliconia latispatha (Heliconiaceae), Monstera acuminata (Araceae), Myriocarpa longipes (Urticaceae), Piper hispidum (Piperaceae), Poulsenia armata (Moraceae), Pseudolmedia oxyphyllaria (Moraceae), Salacia megistophylla (Celastraceae), Siparuna thecaphora (Siparunaceae) and Syngonium podophyllum (Araceae). NDMS ordination and cluster analysis grouped the phorophytes into hierarchically structured clusters variously correlated with microsite, phorophyte species and foliicolous lichen species richness. Indicator species analysis revealed statistically significant foliicolous lichen species characteristic for terminal clusters and for phorophyte species. We conclude that the principle of “diversity begets diversity” may apply, in that phorophyte diversity influences the diversity of foliicolous lichen communities through the manifestation of subtle phorophyte preferences, best seen in well‐developed communities on leaves with higher longevity. Thus, well‐preserved forest ecosystems, with a higher diversity of suitable phorophytes, will support a higher diversity of foliicolous lichens, a phenomenon that extents to epiphytes in general.

It is now a decade since The International Commission on the Taxonomy of Fungi (ICTF) produced an overview of requirements and best practices for describing a new fungal species. In the meantime the International Code of Nomenclature for algae, fungi, and plants (ICNafp) has changed from its former name (the International Code of Botanical Nomenclature) and introduced new formal requirements for valid publication of species scientific names, including the separation of provisions specific to Fungi and organisms treated as fungi in a new Chapter F. Equally transformative have been changes in the data collection, data dissemination, and analytical tools available to mycologists. This paper provides an updated and expanded discussion of current publication requirements along with best practices for the description of new fungal species and publication of new names and for improving accessibility of their associated metadata that have developed over the last 10 years. Additionally, we provide: (1) model papers for different fungal groups and circumstances; (2) a checklist to simplify meeting (i) the requirements of the ICNafp to ensure the effective, valid and legitimate publication of names of new taxa, and (ii) minimally accepted standards for description; and, (3) templates for preparing standardized species descriptions.

The taxonomy of the genus Sticta in Hawaii is reassessed, based on a separately published molecular phylogeny using the fungal barcoding marker ITS. Based on Magnusson and Zahlbruckner's treatment from 1943 and Magnusson's catalogue from 1955, seven species of Sticta and three infraspecific taxa had been reported from the archipelago, all widespread except the putative endemic S. plumbicolor. Here we provide a taxonomic treatment of 13 taxa, 12 species and one subspecies, distinguished in a previous phylogenetic analysis: S. acyphellata, S. andina, S. antoniana, S. emmanueliana, S. flynnii, S. fuliginosa, S. hawaiiensis, S. limbata, S. plumbicolor, S. scabrosa subsp. hawaiiensis, S. smithii, S. tomentosa and S. waikamoi. All taxa are described, discussed and illustrated and a dichotomous key is presented. The implications of revised species taxonomies for studies in other fields such as ecology, ecophysiology, biogeography, biochemistry, and applications such as environmental monitoring are discussed. We also propose a protocol to use Sticta lichens to monitor the environmental health of Hawaiian ecosystems.

The known collections of the genus Cora in continental North America north of Mexico, all restricted to Florida, are shown to belong to a single species, representing a previously unrecognized taxon formally described herein as C. timucua. Based on data of the fungal ITS barcoding marker, obtained through Sanger and Illumina sequencing from two historical collections, the new species is phylogenetically most closely related to C. casanarensis from Colombia and C. itabaiana from Brazil, although it is morphologically most similar to the only distantly related C. hymenocarpa from Costa Rica. Based on data from the Consortium of North American Lichen Herbaria (CNALH) and from the Global Biodiversity Information Facility (GBIF), most of the collections of C. timucua originate from around the turn of the 19th century, while a few were made in the second half of the 20th century, all between 1968 and 1985. Almost all collections originate from Florida sand pine scrub, apparently the preferred habitat of this taxon. Neither modern collections nor extant localities are known. Based on these findings and the substantial degree of land use change in Florida in the past decades, we assessed the conservation status of C. timucua using the IUCN Red List criteria and found that it should be classified as critically endangered (CR), in line with the status of another Florida endemic, Cladonia perforata, which was the first federally red-listed lichen in the United States. The most likely location where C. timucua may still be extant is Ocala National Forest in the north-central portion of the Florida peninsula, although recent macrolichen surveys in that area did not encounter this species.

We present the first broad molecular-phylogenetic revision of the lichenized family Gomphillaceae, based on 408 newly generated sequences of the mitochondrial SSU rDNA and nuclear LSU rDNA, representing 342 OTUs. The phylogenetic analysis of 20 out of the 28 currently accepted genera resulted in 48 clades. Twelve genera were resolved as monophyletic: Actinoplaca, Arthotheliopsis, Bullatina, Caleniopsis, Corticifraga, Gomphillus, Gyalectidium, Gyalidea, Jamesiella, Rolueckia, Rubrotricha, and Taitaia. Two genera resulted paraphyletic, namely Aulaxina (including Caleniopsis) and Asterothyrium (including Linhartia). Six genera were in part highly polyphyletic: Aderkomyces, Calenia, Echinoplaca, Gyalideopsis, Psorotheciopsis, and Tricharia. While ascoma morphology and anatomy has traditionally been considered as main character complex to distinguish genera, our study supported the notion that the characteristic asexual anamorph of Gomphillaceae, the so-called hyphophores, are diagnostic for most of the newly recognized clades. As a result, we recognize 26 new genus-level clades, three of which have names available (Microxyphiomyces, Psathyromyces, Spinomyces) and 23 that will require formal description as new genera. We also tested monophyly for 53 species-level names for which two or more specimens were sequenced: 27 were supported as monophyletic and representing a single species, 13 as monophyletic but with an internal topology suggesting cryptic speciation, four as paraphyletic, and nine as polyphyletic. These data suggest that species richness in the family is higher than indicated by the number of accepted names (currently 425); they also confirm that recently refined species concepts reflect species richness better than the broad concepts applied in Santesson's monograph. A divergence time analysis revealed that foliicolous Gomphillaceae diversified after the K-Pg-boundary and largely during the Miocene, a notion supported by limited data available for other common foliicolous lineages such as Chroodiscus (Graphidaceae), Pilocarpaceae, and Porinaceae. This contradicts recent studies suggesting that only macrofoliose Lecanoromycetes exhibit increased diversification rates in the Cenozoic.

Historical and recent material of Bacidia akompsa (basionym: Biatora akompsa) was studied to determine the taxonomic position of this crustose lichen, prompted by the appearance of the provisional name “Lecanactis akompsa (Tuck.) ined.” in multiple online lichen databases and checklists. Initial morphological assessment of verified material confirmed B. akompsa as a member of Arthoniales but distinct from Lecanactis by having epruinose ascomata with a thin exciple open below the hymenium; a pale, K/I+ violet to deep blue hypothecium; and ascospores with a gelatinous sheath. Molecular phylogenetic analysis of specimens recently collected near the type locality revealed B. akompsa to be nested within the genus Pentagenella in Opegraphaceae. The same placement was found using phenotype-based phylogenetic binning with 37 morphological, anatomical and chemical characters. The new combination Pentagenella akompsa is proposed and the species represents the first North American member of this genus, otherwise known only from Chile and Peru. Therefore, our results reveal the genus Pentagenella to have a remarkably disjunct distribution between North and South America. Several specimens previously labeled as Bacidia, Biatora, or Lecanactis akompsa were found to be misidentifications, narrowing the range of P. akompsa to coastal California, where it is found on two coastal conifer species that are (like P. akompsa itself) narrowly endemic and of conservation concern.

Recent studies have uncovered remarkable diversity in Dictyonema s.lat. basidiolichens, here recognized as subtribe Dictyonemateae. This group includes five genera and 148 species, but hundreds more await description. The photobionts of these lichens belong to Rhizonema, a recently resurrected cyanobacterial genus known by a single species. To further investigate photobiont diversity within Dictyonemateae, we generated 765 new cyanobacterial sequences from 635 specimens collected from 18 countries. The ITS barcoding locus supported the recognition of 200 mycobiont (fungal) species among these samples, but the photobiont diversity was comparatively low. Our analyses revealed three main divisions of Rhizonema, with two repeatedly recovered as monophyletic (proposed as new species), and the third mostly paraphyletic. The paraphyletic lineage corresponds to R. interruptum and partnered with mycobionts from all five genera in Dictyonemateae. There was no evidence of photobiont-mycobiont co-speciation, but one of the monophyletic lineages of Rhizonema appears to partner predominantly with one of the two major clades of Cora (mycobiont) with samples collected largely from the northern Andes. Molecular clock estimations indicate the Rhizonema species are much older than the fungal species in the Dictyonemateae, suggesting that these basidiolichens obtained their photobionts from older ascolichen lineages and the photobiont variation in extant lineages of Dictyonemateae is the result of multiple photobiont switches. These results support the hypothesis of lichens representing "fungal farmers," in which diverse mycobiont lineages associate with a substantially lower diversity of photobionts by sharing those photobionts best suited for the lichen symbiosis among multiple and often unrelated mycobiont lineages.

Symbioses are evolutionarily pervasive and play fundamental roles in structuring ecosystems, yet our understanding of their macroevolutionary origins, persistence, and consequences is incomplete. We traced the macroevolutionary history of symbiotic and phenotypic diversification in an iconic symbiosis, lichens. By inferring the most comprehensive time-scaled phylogeny of lichen-forming fungi (LFF) to date (over 3,300 species), we identified shifts among symbiont classes that broadly coincided with the convergent evolution of phylogenetically or functionally similar associations in diverse lineages (plants, fungi, bacteria). While a relatively recent loss of lichenization in Lecanoromycetes was previously identified, our work instead suggests lichenization was abandoned far earlier, interrupting what had previously been considered a direct switch between trebouxiophycean and trentepohlialean algal symbionts. Consequently, some of the most diverse clades of LFF are instead derived from nonlichenized ancestors and re-evolved lichenization with Trentepohliales algae, a clade that also facilitated lichenization in unrelated lineages of LFF. Furthermore, while symbiont identity and symbiotic phenotype influence the ecology and physiology of lichens, they are not correlated with rates of lineage birth and death, suggesting more complex dynamics underly lichen diversification. Finally, diversification patterns of LFF differed from those of wood-rotting and ectomycorrhizal taxa, likely reflecting contrasts in their fundamental biological properties. Together, our work provides a timeline for the ecological contributions of lichens, and reshapes our understanding of symbiotic persistence in a classic model of symbiosis.

Phylogenetic analysis of some foliicolous lichens collected in Hainan Province, China, revealed a new lineage morphologically similar to Porina but phylogenetically related to Strigulaceae (Dothideomycetes), differing from the latter in ascus type. The monospecific genus Tenuitholiascus gen. nov. is introduced for the single species, T. porinoides sp. nov., which is placed in the new, monogeneric family Tenuitholiascaceae, sister to Strigulaceae in Strigulales. The new taxon closely resembles the genus Porina in external morphology and ascospore type, as well as the thin-walled asci and unbranched paraphyses. Yet, it is entirely unrelated to the latter, which belongs in class Lecanoromycetes in the order Gyalectales.

This paper discusses three issues that challenge contemporaneous taxonomy, with examples from the fields of mycology and lichenology, formulated as three questions: (1) What is the importance of taxonomy in contemporaneous and future science and society? (2) An increasing methodological gap in alpha taxonomy: challenge or opportunity? (3) The Nagoya Protocol: improvement or impediment to the science of taxonomy? The importance of taxonomy in society is illustrated using the example of popular field guides and digital media, a billion-dollar business, arguing that the desire to name species is an intrinsic feature of the cognitive component of nature connectedness of humans. While continuous societal support of a critical mass of taxonomists is necessary to catalogue all species on Earth, it is shown that this is a finite task, and a proposal is made how a remaining 10 million species can be catalogued within 40 years by 1,000 well-trained and dedicated taxonomists, with an investment of $4 billion, corresponding to 0.0001% of the annual global GDP or 0.005% of annual military expenditures. Notorious undercitation of actually used taxonomic resources and lack of coverage of impact metrics for monographs and other taxonomic work that cannot be published in indexed journals is discussed and suggestions are made how this problem can be remedied. An increasing methodological gap in approaches to taxonomy, between classic morphological and advanced genomic studies, affects in particular taxonomists in biodiversity-rich countries and amateurs, also regarding proper training to apply advanced methods and concepts. To counterbalance this problem, international collaborations bringing different expertise to the table and undertaking mutual capacitation are one successful remedy. Classic taxonomy still works for many groups and is a first approach to catalogue species and establish taxon hypotheses, but ultimately each taxonomic group needs to be studied with the array of methods proper to the group, including descriptive work. Finally, the Convention on Biological Diversity (CBD) and the Nagoya Protocol has put additional burden on basic biodiversity science. Using lichenology in Latin America and Brazil as an example, it is shown that the spirit of non-monetary benefit-sharing proper to taxonomy and systematics, namely capacitation, joint publications, and shared reference collections, has been increasingly implemented long before the CBD and the Nagoya Protocol, and does not need additional “policing”. Indeed, the Nagoya Protocol puts the heaviest burden on taxonomy and researchers cataloguing biodiversity, whereas for the intended target group, namely those seeking revenue gain from nature, the protocol may not actually work effectively. The notion of currently freely accessible digital sequence information (DSI) to become subject to the protocol, even after previous publication, is misguided and conflicts with the guidelines for ethical scientific conduct. Through its implementation of the Nagoya Protocol, Colombia has set a welcome precedence how to exempt taxonomic and systematic research from “access to genetic resources”, and hopefully other biodiversity-rich countries will follow this example.

DNA barcoding using the nuclear internal transcribed spacer (ITS) has become prevalent in surveys of fungal diversity. This approach is, however, associated with numerous caveats, including the desire for speed, rather than accuracy, through the use of automated analytical pipelines, and the shortcomings of reference sequence repositories. Here we use the case of a specimen of the bracket fungus Trametes s.lat. (which includes the common and widespread turkey tail, T. versicolor) to illustrate these problems. The material was collected in Vietnam as part of a biodiversity inventory including DNA barcoding approaches for arthropods, plants and fungi. The ITS barcoding sequence of the query taxon was compared against reference sequences in GenBank and the curated fungal ITS database UNITE, using BLASTn and MegaBLAST, and was subsequently analysed in a multiple alignment-based phylogenetic context through a maximum likelihood tree including related sequences. Our results initially indicated issues with BLAST searches, including the use of pairwise local alignments and sorting through Total score and E value, rather than Percentage identity, as major shortcomings of the DNA barcoding approach. However, after thorough analysis of the results, we concluded that the single most important problem of this approach was incorrect sequence labelling, calling for the implementation of third-party annotations or analogous approaches in primary sequence repositories. In addition, this particular example revealed problems of improper fungal nomenclature, which required reinstatement of the genus name Cubamyces (= Leiotrametes), with three new combinations: C. flavidus, C. lactineus and C. menziesii. The latter was revealed as the correct identification of the query taxon, although the name did not appear among the best BLAST hits. While the best BLAST hits did correspond to the target taxon in terms of sequence data, their label names were misleading or unresolved, including [Fungal endophyte], [Uncultured fungus], Basidiomycota, Trametes cf. cubensis, Lenzites elegans and Geotrichum candidum (an unrelated ascomycetous contaminant). Our study demonstrates that accurate identification of fungi through molecular barcoding is currently not a fast-track approach that can be achieved through automated pipelines.Citation: Lucking R., Truong B. V., Huong D. T. T., Le N. H., Nguyen Q. D., Nguyen V. D., Raab-Straube E. von, Bollendorff S., Govers K. & Di Vincenzo V. 2020: Caveats of fungal barcoding: a case study in Trametes s.lat. (Basidiomycota: Polyporales) in Vietnam reveals multiple issues with mislabelled reference sequences and calls for third-party annotations. – Willdenowia 50: 383–403. doi: https://doi.org/10.3372/wi.50.50302Version of record first published online on 15 September 2020 ahead of inclusion in December 2020 issue.

Strigulasensu lato has been previously defined based on phenotype characters as a rather broad genus including tropical to temperate species growing on a wide array of substrata. In this study, based on a multilocus phylogenetic approach, we show that foliicolous species form six well-delimited clades that correlate with diagnostic phenotype features, including thallus morphology, carbonization of the involucrellum and excipulum, ascospore dimensions, and type of macroconidia. Given the topology, with five of the six clades emerging on long stem branches, and the strong phenotypical differentiation between the clades, we recognize these at the genus level, making mostly use of previously established genus names. Four genera, namelyPhylloporis,Puiggariella,Raciborskiella, andRacoplaca, are resurrected for theS. phyllogena,S. nemathora,S. janeirensis, andS. subtilissimagroups, respectively, whereas one new genus,Serusiauxiellagen. nov., is introduced for a novel lineage with peculiar macroconidia. The only sequenced non-foliicolous species,S. jamesii, is not closely related to these six foliicolous lineages but clusters withFlavobatheliumandPhyllobathelium, revealing it as an additional undescribed genus-level lineage being treated elsewhere. Within the new genusSerusiauxiella, three new species are described:Serusiauxiella filiferasp. nov.,S. flagellatasp. nov., andS. sinensissp. nov. In addition, ten new combinations are proposed:Phylloporis austropunctatacomb. nov.,P. radiatacomb. nov.,P. vulgariscomb. nov.,Puiggariella confluenscomb. et stat. nov.,P. nemathoracomb. nov.,P. nigrocinctacomb. nov.,Racoplaca maculatacomb. nov.,R. melanobaphacomb. nov.,R. transversoundulata, andR. tremenscomb. nov. We also report on a peculiar, previously unrecognized growth behaviour of the macroconidial appendages inStrigulas.lat.

The genusCoenogoniumEhrenb. in Tasmania comprises seven species. New to science are:C. atherospermatisKantvilas, Rivas Plata & Lücking, endemic to Tasmania and characterized by pale yellowish beige apothecia and relatively small ascospores, 6–8·5×2·5–3 µm;C. urceolatumKantvilas, Rivas Plata & Lücking, likewise endemic to Tasmania and characterized by orange, urceolate apothecia, 0·3–0·4 mm wide, and uniseriate ascospores, 8·5–11×2·5–3 µm; andC. australienseKantvilas & Lücking, recorded from Tasmania, South Australia and New South Wales, and characterized by orange apothecia, 0·5–2 mm wide, and relatively broad ascospores, 10–14×3–4·5 µm. Also treated areC. lutescens(Vĕzda & Malcolm) Malcolm (Tasmania and New Zealand) and three widespread taxa, namelyC. implexumNyl. (Southern Hemisphere),C. luteum(Dicks.) Kalb & Lücking andC. pineti(Schrad. ex Ach.) Lücking & Lumbsch (both subcosmopolitan). All species are described in full from Tasmanian collections and illustrated, and their ecology, variation and affinities to related species are discussed. The Tasmanian taxa are also discussed in the context of the Australasian lichen biota.

Nuclear ribosomal internal transcribed spacer (ITS) region is well-established as universal DNA barcode marker for Fungi. Here we examined the utility of DNA barcoding for delimiting species in lichenized Basidiomycota from Colombia, focusing on the genus Sulzbacheromyces (Lepidostromatales). The topology of the best-scoring maximum likelihood tree based on ITS data shows eight separate, well-supported lineages within Sulzbacheromyces, including the six already known species S. caatingae from Brazil, S. miomboensis De Kesel & Ertz from Africa, and S. bicolor, S. fossicola, S. sinensis, and S. yunnanensis from Asia. In addition, two further, sympatric and semi-cryptic species from the Choco Biogeographic Region, one of the most diverse regions of the world, are recognized: S. chocoensis Coca, Lucking & Moncada sp. nov and S. tutunendo Coca, Lucking & Moncada sp. nov. Both form separate, strongly supported species-level lineages, but differ morphologically from each other and from S. caatingae in minor details only, which are best assessed with fresh material in the field.

Os objetivos deste estudo são sumarizar e apresentar uma listagem de espécies de fungos conidiais e liquenizados da Floresta Nacional (FLONA) de Caxiuanã, localizada no estado do Pará, Brasil. Os dados desta listagem foram obtidos com base em coleções de herbário e na literatura, ambos resultantes de expedições realizadas nesta FLONA entre 1995 e 2013. Na maioria, estes fungos pertencem ao filo Ascomycota. Um total de 233 espécies foi identificado paraCaxiuanã, sendo 56 de fungos conidiais e 177 de fungos liquenizados. As espécies Septomyrothecium uniseptatum Matsush. e Pyrenidium zamiae (Müll. Arg.) Matzer representam os primeiros registros para o Brasil. Novos registros para o Pará e para a FLONA de Caxiuanã são apresentados. A distribuição geográfica no Brasil é referida para todas as espécies. Estes resultados demonstram a importância da realização de estudos sobre a micobiota de áreas que apresentem dados escassos ou inexistentes.

Quercus humboldtii es un elemento dominante en los bosques altoandinos de Boyaca, Colombia, el cual, a pesar de ser una especie sombrilla, registra consumos per-capita de 1300 Kg.ano -1 y el remplazo de sus rodales por plantaciones de Pinus patula . Este remplazo afecta las comunidades liquenicas mas sensibles. Para rastrear dichas afecciones se examinaron los liquenes en cuatro arboles de cada especie forofita. Cada arbol fue dividido en cinco estratos verticales y se midio la rugosidad, humedad, acidez y luminosidad en cada estrato. Los analisis comprendieron indices de diversidad, tablas Anova y escalas multidimensionales. Se registraron 161 especies de liquenes, se reconocen 53 especies exclusivas sobre pino y 45 exclusivas de roble. Se destacan Cladonia ceratophylla y Herpothallon rubrocinctum como especies comunes entre forofitas, y la exclusividad de Astrothelium spp, Anzia leucobatoides y Leptogium diaphanum sobre los robles. La luminosidad y la rugosidad son factores de influencia en la distribucion de los liquenes.

There is urgent need for a formal nomenclature of sequence-based, voucherless Fungi, given that environmental sequencing has accumulated more than one billion fungal ITS reads in the Sequence Read Archive, about 1,000 times as many as fungal ITS sequences in GenBank. These unnamed Fungi could help to bridge the gap between 115,000 to 140,000 currently accepted and 2.2 to 3.8 million predicted species, a gap that cannot realistically be filled using specimen or culture-based inventories. The Code never aimed at placing restrictions on the nature of characters chosen for taxonomy, and the requirement for physical types is now becoming a constraint on the advancement of science. We elaborate on the promises and pitfalls of sequence-based nomenclature and provide potential solutions to major concerns of the mycological community. Types of sequence-based taxa, which by default lack a physical specimen or culture, could be designated in four alternative ways: (1) the underlying sample ('bag' type), (2) the DNA extract, (3) fluorescent in situ hybridization (FISH), or (4) the type sequence itself. Only (4) would require changes to the Code and the latter would be the most straightforward approach, complying with three of the five principal functions of types better than physical specimens. A fifth way, representation of the sequence in an illustration, has been ruled as unacceptable in the Code. Potential flaws in sequence data are analogous to flaws in physical types, and artifacts are manageable if a stringent analytical approach is applied. Conceptual errors such as homoplasy, intragenomic variation, gene duplication, hybridization, and horizontal gene transfer, apply to all molecular approaches and cannot be used as a specific argument against sequence-based nomenclature. The potential impact of these phenomena is manageable, as phylogenetic species delimitation has worked satisfactorily in Fungi. The most serious shortcoming of sequence-based nomenclature is the likelihood of parallel classifications, either by describing taxa that already have names based on physical types, or by using different markers to delimit species within the same lineage. The probability of inadvertently establishing sequence-based species that have names available is between 20.4 % and 1.5 % depending on the number of globally predicted fungal species. This compares favourably to a historical error rate of about 30 % based on physical types, and this rate could be reduced to practically zero by adding specific provisions to this approach in the Code. To avoid parallel classifications based on different markers, sequence-based nomenclature should be limited to a single marker, preferably the fungal ITS barcoding marker; this is possible since sequence-based nomenclature does not aim at accurate species delimitation but at naming lineages to generate a reference database, independent of whether these lineages represent species, closely related species complexes, or infraspecies. We argue that clustering methods are inappropriate for sequence-based nomenclature; this approach must instead use phylogenetic methods based on multiple alignments, combined with quantitative species recognition methods. We outline strategies to obtain higher-level phylogenies for ITS-based, voucherless species, including phylogenetic binning, 'hijacking' species delimitation methods, and temporal banding. We conclude that voucherless, sequence-based nomenclature is not a threat to specimen and culture-based fungal taxonomy, but a complementary approach capable of substantially closing the gap between known and predicted fungal diversity, an approach that requires careful work and high skill levels.

Seven proposals to modify the provisions of the International Code of Nomenclature for algae, fungi, and plants (ICN) at the 11th International Mycological Congress (IMC11) in July 2018 had been received by the proscribed date of 1 March 2018. These proposals are formally presented together here. The topics addressed relate to the clarification of the meaning of “original material” in relation to the typification of sanctioned names, the indication of the nomenclatural status of sanctioned names in author citations, the use of DNA sequences as nomenclatural types, and the possibility of including repository identifiers as an alternative to author citations. A synopsis of the proposals will be provided during April, and the mycological community will be invited to provide a guiding vote up to 10 June 2018. Final decisions on these proposals are to be made following debate at the Fungal Nomenclature Session of IMC11.

21 new records of lichens from the family Graphidoideae (Graphidaceae) for Colombia and seven for the state of Valle del Cauca are presented. These new reports correspond to the genera Ampliotrema, Chapsa, Gyrotrema, Myriotrema, Ocellularia, Stegobolus, Thelotrema, and Wirthiotrema. These records expand the list of known species for the country of Ampliotrema to 5, Chapsa to 14, Gyrotrema to 3, Myriotrema to 11, Ocellularia to 29, Stegobolus to 7, and to 2. All these species are conservation indicators for tropical humid forests, for they are found only in low elevation humid forests that are in very good state.

We present a phylogenetic revision of theSticta filixmorphodeme in New Zealand. This non-monophyletic group of early diverging clades in the genusStictais characterized by a stalked thallus with a green primary photobiont and the frequent formation of a dendriscocauloid cyanomorph. Traditionally, three species have been distinguished in New Zealand:S. filix(Sw.) Nyl.,S. lacera(Hook. f. & Taylor) Müll. Arg. andS. latifronsA. Rich., with two cyanomorphs separated under the namesDendriscocaulon dendriothamnodesDughi ex D. J. Galloway (traditionally associated withS. latifrons) andD. dendroides(Nyl.) R. Sant. ex H. Magn. (traditionally associated withS. filix).Sticta lacerawas not included in the present study due to the lack of authentic material (all specimens originally identified under that name and sequenced clustered withS. filix);S. filixwas confirmed as a distinct species whereasS. latifronss. lat. was shown to represent two unrelated species,S. latifronss. str. and the reinstatedS. menziesiiHook. f. & Taylor. The cyanomorphs ofS. filixandS. latifronsare not conspecific with the types of the namesD. dendriothamnodesandD. dendroides, respectively; theD. dendriothamnodescyanomorph belongs to the Australian taxonSticta stipitataC. Knight ex F. Wilson, which is not present in New Zealand, whereas theD. dendroidescyanomorph corresponds to a previously unrecognized species with unknown chloromorph, recombined here asSticta dendroides(Nyl.) Moncada, Lücking & de Lange. Thus, instead of three species (S. filix,S. lacera,S. latifrons) with their corresponding cyanomorphs, five species are now distinguished in this guild in New Zealand:S. dendroides(cyanomorph only),S. filix(chloro- and cyanomorph),S. lacera(chloromorph only),S. latifrons(chloro- and cyanomorph) andS. menziesii(chloro- and cyanomorph). A key is presented for identification of the chloromorphs and the dendriscocauloid cyanomorphs of all species. Semi-quantitative analysis suggests that species in this guild are good indicators of intact forest ecosystems in New Zealand and that the two newly recognized species,S. dendroidesandS. menziesii, appear to perform particularly well in this respect. The use of lichens as bioindicators of environmental health is not yet established in New Zealand and so, based on our results, we make the case to develop this approach more thoroughly.

We assessed the taxonomy and phylogenetic relationships of Hawaiian species of the neotropical genusLobariella. A single species was previously reported from the archipelago, the widespread neotropicalL. 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 thatL. crenulatas. str. is seemingly present in the archipelago based on older herbarium collections. However, HawaiianLobariellaincludes three additional, presumably endemic, species unrelated toL. crenulataand new to science:L. flynnianaLücking, Moncada & C. W. Sm., with richly branched marginal phyllidia giving the thallus a fruticose appearance;L. robustaLücking, Moncada & C. W. Sm., with a rather thick thallus and abundant, large, laminal phyllidia; andL. sandwicensisLücking, Moncada & C. W. Sm., with a delicate thallus producing apothecia.Lobariella flynnianarepresents 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.

Molecular studies have shown the type collection of Omphalina oreades to be conspecific with a small brown basidiolichen from the Appalachian range in Newfoundland, both with 4-spored basidia. Two sequences deposited in GenBank, originally identified as O. grisella, fell in the same clade. Sequences of the type collection of Omphalia grisella, with 2-spored basidia, formed a sister clade together with two GenBank deposits, one identified as O. grisella and the other as Omphalina velutina. Omphalina oreades is recombined here as Lichenomphalia oreades comb. nov., and the species redescribed and illustrated. Sequences of the internal transcribed spacer regions of nuclear ribosomal DNA (ITS rDNA) from the algae associated with two collections of L. oreades fell within a highly supported clade with members of an undetermined species of Coccomyxa. The most abundant algal ribosomal large subunit sequence from granules at the base of a different collection matched GenBank sequences identified as Chloroidium ellipsoideum, which is probably either a free-living algal species or a partner to a species of Trapeliopsis. The second most abundant sequence matched Coccomyxa subellipsoidea and is the most likely photobiont of L. oreades. Further studies are required to elucidate the relationship of L. velutina to these taxa.

Ramalina europaeaGasparyan, Sipman & Lücking andR. labiosorediataGasparyan, Sipman & Lücking, two species of theR. pollinariagroup, are described here as new to science.Ramalina europaea, widely distributed in Europe, can be distinguished by small, punctiform, often terminal soralia starting out on small, spine-like branchlets, whereasR. labiosorediatafrom North America differs fromR. pollinarias. str. andR. europaeain the almost exclusively terminal soralia formed on the tips of normal lobes, originating from the underside and becoming irregularly labriform. Morphological characters, chemistry, ecology and geographical distribution are discussed and a key to the species of theRamalina pollinariagroup is provided. The topology of a maximum likelihood tree based on ITS shows the presence of three well-supported clades, corresponding to the morphological differences of the three species. The status of several historical names variously placed in synonymy with or described as infraspecific entities ofR. pollinariais reassessed and a new neotype and an epitype are designated forLichen pollinarius, a neotype forL. squarrosus, making it a synonym ofR. farinacea, and lectotypes forR. pollinariavar.elatior, making it a synonym ofR. pollinarias. str., and for var.humilis, a taxon of yet unknown affinity.

Diederich, P., Lucking, R., Aptroot, A., Sipman, H. J. M., Braun, U., Ahti, T. & Ertz, D. 2017. New species and new records of lichens and lichenicolous fungi from the Seychelles. — Herzogia 30: 182–236. Sixteen species of lichens and lichenicolous fungi from the Seychelles are described as new: Abrothallus ramalinae (on Ramalina), Coenogonium beaverae, Fissurina seychellensis, Fulvophyton macrosporum, Graphis lindsayana, Nigrovothelium inspersotropicum, Opegrapha salmonea, Porina morelii, Pseudopyrenula media, Ramichloridium tropicum (on sterile lichen with Trentepohlia), Sarcographa praslinensis, S. subglobosa, Stictographa dirinariicola (on Dirinaria picta), Stirtonia epiphylla, Talpapellis mahensis (on sterile lichen with Trentepohlia) and Trimmatothele petri; Abrothallus ramalinae is also reported from Australia, New Zealand and Papua New Guinea, and Nigrovothelium inspersotropicum from Guyana. The following 29 species are new to Africa: Acanthothecis asprocarpa, Amandinea diorista var. hyp...

Two species of lichen-forming fungi with pyrenocarpous ascomata are described as new to science: Aspidothelium silverstonei and Astrothelium fuscosporum. The epiphytic lichens were found in Choco Biogeographic region of Colombia. Aspidothelium silverstonei is characterized by the largest muriform ascospores known in the genus, while Astrothelium fuscosporum is the first species of Astrothelium known to produce pigmented ascospores.

The history of assigning ranks to fungi, as well as the relative importance of using divergence time estimates is reviewed. The paper pays tribute to the major mycological players, and especially to David Hawksworth on his 70th birthday and his contribution to fungal ranking in Systema Ascomycetum from 1982 to 1998. Following the conclusion of the latter series, the ranking continued with the Outlines of Ascomycota in 2007 and 2010 and more recently with specific classes in 'Towards an outline of Sordariomycetes' and 'Families of Dothideomycetes'. Earlier classifications based on phenotype were certainly more subjective; however, remarkably many of these old arrangements have stood the test of time. More recently, phylogenetic analyses have provided evidence towards a natural classification, resulting in significant changes in many lineages. The classification arrangements however, are still subjective and dependent on the taxa analysed, resulting in different taxonomic interpretations and schemes, particularly when it comes to ranking. Thus, what have been considered as genera by some, have been introduced as families by others. More recently, estimation of divergence times using molecular clock methods have been used as objective evidence for higher ranking of taxa. A divergence period (i.e. 200-300 MYA) can be used as a criterion to infer when a group of related taxa evolved and what rank they should be given. We compiled data on divergence times for various higher ranking taxa in the Kingdom Fungi. The kingdom evolved 1000-1600 MYA (Stenian-Calymmian), while the presently accepted phyla evolved between 358 and 541 MYA (Devonian-Cambrian). Divergence times for subphyla are generally between 358 and 485 MYA (Devonian-Ordovician), those of classes 145-358 MYA (Jurassic-Carboniferous), subclasses 66-358 MYA (Cretaceous-Carboniferous), orders 23-252 MYA (Paleogene-Triassic), families 2.8-145 MYA (Neogene-Cretaceous), and genera 2.8-66 MYA (Neogene-Paleogene). Thus, there are wide discrepancies in the times different taxa diverged. We provide an overview over Ascomycota, showing how application of temporal banding could affect the recognition of higher taxa at certain rank levels. We then use Sordariomycetes as an example where we use divergence times to provide additional evidence to stabilize ranking of taxa below class level. We propose a series of evolutionary periods that could be used as a guide to determine the various higher ranks of fungi: phyla[550 MYA, subphyla 400-550 MYA; classes 300-400 MYA; subclasses 250-300 MYA, orders 150-250 MYA, and families 50-150 MYA. It is proposed that classification schemes and ranking of taxa should, where possible, incorporate a polyphasic approach including phylogeny, phenotype, and estimate of divergence times.

Based on an unexpected result of obtaining molecular sequence data from tropical representatives of the genus Normandina, we revised the biological concept of the neotropical taxon Marchandiomphalina foliacea. The obtained data let us conclude that M. foliacea is not a basidiomycete, as originally proposed, but belongs in Verrucariaceae, in the genus Agonimia, including its perithecia which had been identified with the lichenicolous Norrlinia peltigericola. The ITS (and nuLSU) sequences previously obtained from M. foliacea, seemingly confirming its status as a basidiomycete, are from an unmanifested lichenicolous fungus, present also in numerous specimens of Normandina. ITS data suggest the presence of seven lineages that can be recognized at the species level, forming two clusters: one cluster of three lineages found in thalli of M. foliacea, and a second cluster of four lineages found in thalli of Normandina. This pattern is similar to what has recently been found in the basidiomycete genus Cyphobasidium occurring predominantly in Parmeliaceae lichens. We propose the combination of Omphalina foliacea into the genus Agonimia, as Agonimia foliacea (P.M. Jorg.) Lucking & Moncada, comb. nov., and place Marchandiomphalina in synonymy with Agonimia. To formally recognize the unnamed lichenicolous basidiomycete present in Agonimia and Normandina thalli, we take advantage of provision ICN Art. 40.5 in the Code and describe the unmanifested fungus as a new genus, with seven new species, even if no physical type specimens can be preserved (except for the corresponding host lichens which, however, do not show the features of the fungus): Lawreymyces Lucking & Moncada, gen. nov. (Type: L. palicei), with L. bogotensis Lucking & Moncada, sp. nov., L. columbiensis Lucking & Moncada, sp. nov., L. confusus Lucking & Moncada, sp. nov., L, foliaceae Lucking & Moncada, sp. nov., L. palicei Lucking & Moncada, sp. nov., L. pulchellae Lucking & Moncada, sp. nov., and L. spribillei Lucking & Moncada, sp. nov. This opens the door to the formal recognition of thousands of species of voucherless fungi detected through environmental sequencing techniques under the current Code.

This study is a re-assessment of basidiolichen diversity in the Galapagos Islands. We present a molecular phylogenetic analysis, based on 92 specimens from Galapagos, using two nuclear ribosomal DNA markers (ITS and nuLSU). We also re-examined the morphology and anatomy of all sequenced material. The molecular results confirm our previous assessment that all Galapagos basidiolichens belong to the Dictyonema clade, which in Galapagos is represented by four genera: Acantholichen, Cora, Cyphellostereum, and Dictyonema. Most species previously reported from Galapagos in these genera were at the time believed to represent widely distributed taxa. This conclusion, however, has changed with the inclusion of molecular data. Although almost the same number of species is distinguished, the phylogenetic data now suggest that all are restricted to the Galapagos Islands. Among them, six species are proposed here as new to science, namely Cora galapagoensis, Cyphellostereum unoquinoum, Dictyonema barbatum, D. darwinianum, D. ramificans, and D. subobscuratum; and four species have already been described previously, namely Acantholichen galapagoensis, Cora santacruzensis, Dictyonema pectinatum, and D. galapagoense, here recombined as Cyphellostereum galapagoense. Our analysis is set on a very broad phylogenetic framework, which includes a large number of specimens (N = 826) mainly from Central and South America, and therefore strongly suggests an unusually high level of endemism previously not recognized. This analysis also shows that the closest relatives of half of the basidiolichens now found in Galapagos are from mainland Ecuador, implying that they reached the islands through the shortest route, with all species arriving on the islands through independent colonization events.

Phylogenetic studies of the lichenized fungal family Graphidaceae necessitate various nomenclatural changes. Tribe Wirthiotremateae is more narrowly circumscribed on the basis of new molecular data and phenotypical differences towards allied clades. The new genus Austrotrema is described to accommodate the three species of the Thelotrema bicinctulum group, Asteristion is resurrected for the seven species of the Chapsa platycarpa group, and Nadvornikia is expanded to include two non-mazaediate species in addition to its two mazaediate species. Asteristion australianum is newly described for Australian material previously identified as T. albo-olivaceum. Eleven new combinations are made: Asteristion alboannuliforme (Bas.: Thelotrema alboannuliforme), As. albo-olivaceum (Bas.: T. albo-olivaceum), As. cupulare (Bas.: T. cupulare), As. leucophthalmum (Bas.: T. leucophthalmum), As. platycarpoides (Bas.: T. platycarpoides), As. platycarpum (Bas.: T. platycarpum), Austrotrema bicinctulum (Bas.: T. bicinc...

Lucking, R., Mangold, A. & Lumbsch, H. T. 2016. A worldwide key to species of the genera Myriotrema and Glaucotrema (lichenized Ascomycota: Graphidaceae), with a nomenclatural checklist of species published in Myriotrema. — Herzogia 29: 493–513. A worldwide key to 54 species currently accepted in the genera Myriotrema sensu Frisch (50) and Glaucotrema (4) is presented, including six species with provisional placement in Myriotrema. We also provide a nomenclatural checklist of all 138 names published in Myriotrema and their current status, with the following eleven new combinations being proposed: Myriotrema leucohymenium, M. zollingeri, Ocellularia craterella, O. extendens, O. masonhalei, O. parvidisca, O. pertusarioides, O. squamuloides, Schizotrema flavolucens and Thelotrema configuratum.

Phylogenetic analysis of some foliicolous lichens collected in Hainan Province, China, revealed a new lineage morphologically similar to Porina but phylogenetically related to Strigulaceae (Dothideomycetes), differing from the latter in ascus type. The monospecific genus Tenuitholiascus gen. nov. is introduced for the single species, T. porinoides sp. nov., which is placed in the new, monogeneric family Tenuitholiascaceae, sister to Strigulaceae in Strigulales. The new taxon closely resembles the genus Porina in external morphology and ascospore type, as well as the thin-walled asci and unbranched paraphyses. Yet, it is entirely unrelated to the latter, which belongs in class Lecanoromycetes in the order Gyalectales.

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Based on separately obtained and analyzed molecular data and within the framework of a global revision of the family Trypetheliaceae, 21 new species are described, from the Neotropics and tropical Asia, in the genera Architrypethelium (1), Astrothelium (15), Bathelium (1), Nigrovothelium (1), Trypethelium (1), and Viridothelium (2), namely: Architrypethelium lauropaluanum Lücking, M. P. Nelsen & Marcelli sp. nov., differing from A. hyalinum in the perithecia immersed between coarse thallus verrucae and in the additional ascospore septa; Astrothelium aurantiacocinereum Lücking, Naksuwankul & Lumbsch sp. nov., differing from A. aeneum in the prominent, well-delimited, trypethelioid pseudostromata and the absence of pigment on the thallus surface, as well as in the barely lichenized thallus; A. carassense Lücking, M. P. Nelsen & Marcelli sp. nov., differing from A. purpurascens in orange, K+ red pseudostroma pigment and the slightly larger ascospores; A. cryptolucens Lücking, M. P. Nelsen & N. Salazar sp. nov., differing from A. carrascoense in the inspersed hymenium; A. fijiense Lücking, Naksuwankul & Lumbsch sp. nov., differing from A. cinereorosellum in the presence of lichexanthone on the well-delimited pseudostromata and in the slightly shorter ascospores; A. laevithallinum Lücking, M. P. Nelsen & Marcelli sp. nov., differing from A. endochryseum in the smooth thallus; A. leucosessile Lücking, M. P. Nelsen & Aptroot sp. nov., differing from A. phlyctaena in the conspicuous, sessile pseudostromata; A. macrostomoides Lücking, M. P. Nelsen & Benatti sp. nov., differing from A. macrostomum in the larger ascospores; A. megacrypticum Lücking, M. P. Nelsen & N. Salazar sp. nov., differing from A. longisporum in the single-spored asci and larger ascospores; A. nicaraguense Lücking, M. P. Nelsen & T. Orozco sp. nov., differing from A. gigantosporum in the smaller ascospores; A. norisianum Lücking, M. P. Nelsen & Aptroot sp. nov., differing from A. sepultum in the distinct, well-delimited pseudostromata; A. obtectum Lücking, M. P. Nelsen & Benatti sp. nov., differing from A. nigrocacuminum in the smaller ascospores; A. sordithecium Lücking, M. P. Nelsen & Marcelli sp. nov., differing from A. leucothelium in the inspersed hymenium and the absence of lichexanthone from the thallus surface outside the pseudostromata; A. subendochryseum Lücking, M. P. Nelsen & Marcelli sp. nov., differing from A. endochryseum in the absence of pigment in the pseudostromata and the lateral thallus cover of the pseudostromata; A. subinterjectum Lücking, M. P. Nelsen & Jungbluth sp. nov., differing from A. obtectum in the smaller pseudostromata and smaller ascospores, and from A. interjectum in the diffuse pseudostromata and smaller ascospores; Bathelium porinosporum Lücking, M. P. Nelsen & Gueidan sp. nov., differing from other Bathelium species in the 3-septate, euseptate ascospores; Nigrovothelium bullatum Lücking, Upreti & Lumbsch sp. nov., differing from N. tropicum in the bullate thallus; Trypethelium tolimense Lücking, Moncada & M. Gut. sp. nov., differing from T. xanthoplatystomum in the absence of a yellow-orange pigment on the pseudostromata and the K+ yellow (not K+ red) medullary pigment; Viridothelium tricolor Lücking, M. P. Nelsen & N. Salazar sp. nov., characterized by black perithecia with a lateral ostiole immersed in white pseudostromata strongly contrasting with the surrounding brown thallus, in combination with 2-spored asci and large, muriform ascospores; and V. vonkonratii Lücking, Naksuwankul & Lumbsch sp. nov., differing from V. virens in larger ascospores and mostly solitary ascomata. All species are illustrated and their taxonomy and phylogenetic relationships are discussed. ITS barcoding sequences are reported for five specimens of Bathelium porinosporum.

A quantitative prediction of global species richness in the lichenized familyTrypetheliaceaeemploying a grid method previously used in similar approaches for the familyGraphidaceaeand the genusCora(Hygrophoraceae) is provided. 421 currently known taxa were used as a base for the calculation and we predict a total of nearly 800 species, corresponding to an increase of almost 100%. Most of the unrecognized taxa are predicted to occur in the Neotropics, which have the highest known species richness. The Palaeotropics are less diverse. In contrast toGraphidaceae, which are mostly confined to rainforests,Trypetheliaceaeshow substantial species richness in dry forest and savannah ecosystems, and future collection efforts should take well-preserved areas representing such ecosystems into consideration.

We provide an expanded and updated, 2-locus phylogeny (mtSSU, nuLSU) of the lichenized fungal family Trypetheliaceae, with a total of 196 ingroup OTUs, in order to further refine generic delimitations and species concepts in this family. As a result, the following 15 clades are recognized as separate genera, including five newly established genera: Aptrootia, Architrypethelium, Astrothelium (including the bulk of corticate species with astrothelioid ascospores; synonyms: Campylothelium, Cryptothelium, Laurera), Bathelium s. str. (excluding B. degenerans and relatives which fall into Astrothelium), the reinstated Bogoriella (for tropical, lichenized species previously placed in Mycomicrothelia), Constrictolumina gen. nov. (for tropical, lichenized species of Arthopyrenia), Dictyomeridium gen. nov. (for a subgroup of species with muriform ascospores previously placed in Polymeridium), Julella (provisionally, as the type species remains unsequenced), Marcelaria (Laurera purpurina complex), Nigrovothelium gen. nov. (for the Trypethelium tropicum group), Novomicrothelia gen. nov. (for an additional species previously placed in Mycomicrothelia), Polymeridium s. str., Pseudopyrenula, Trypethelium s. str. (T. eluteriae group), and Viridothelium gen. nov. (for the Trypethelium virens group). All recognized genera are phenotypically characterized and a discussion on the evolution of phenotypic features in the family is given. Based on the obtained phylogeny, species delimitations are revised and the importance of characters such as thallus morphology, hymenial inspersion, and secondary chemistry for taxonomic purposes is discussed, resulting in a refined species concept.