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The new genus Saxiloba is described with the two species S. firmula from the Caribbean and S. hawaiiensis from Hawaii. Saxiloba is characterized by a unique, placodioid thallus forming distinct lobes, growing on rock in shaded to exposed situations with a trentepohlioid photobiont and a fenestrate thallus anatomy with distinct surface lines. The material is often sterile, but Porina-like perithecia and ascospores had previously been described for the Caribbean taxon and were here confirmed for both species. Molecular sequence data also confirmed placement of this lineage in Porinaceae. Its position within that family supports the notion that Porinaceae should be subdivided into a larger number of genera than proposed in previous classification attempts. Compared to other Porinaceae, Saxiloba exhibits a unique morphology and anatomy that recalls taxa in the related family Graphidaceae and it substantially expands the known phenotypic variation within Porinaceae. The two recognized species are similar in overall morphology but, apart from their disjunct distribution and different substrate ecology, differ in lobe configuration, color and disposition of the crystal clusters and resulting surface patterns.

Foliicolous lichens grow on the surface of living leaves in tropical and subtropical forests. Compared to a large number of species of foliicolous lichens reported from other regions in tropical and subtropical Asia, only six species of the otherwise abundant genus Strigula have been registered from South Korea so far. Three of these, morphologically identified as S. concreta, S. macrocarpa, and S. smaragdula, had previously been shown to share near-identical ITS sequences, casting doubt about the usefulness of this marker for species delimitation in the genus Strigula. To shed light on this conundrum, we surveyed the diversity of the genus Strigula in the Gotjawal forest area on Jeju Island south of mainland Korea, where the climate and vegetation are suitable for foliicolous lichens. As the result of a combined analysis of phenotype and molecular data of the ITS fungal barcoding marker, we found that material morphologically similar to known species formed two strongly supported clades, representing two species new to science, S. depressa Woo, Lücking & Hur sp. nov. and S. multiformis Woo, Lücking & Hur sp. nov., which are described herein. Strigula multiformis included the four previously sequenced specimens identified as S. concreta, S. macrocarpa, and S. smaragdula. A detailed analysis of morphological and anatomical characters revealed that all specimens of S. multiformis were anatomically uniform but varied in thallus morphology, mostly resembling S. smaragdula but with some forms similar to S. concreta and S. macrocarpa, explaining the previous misidentifications. This variation was found to be driven by leaf characters of the phorophyte species, as these apparently influence the morphology of the subcuticular thalli.

New collections are reported of the monospecific genusPolypyrenula, an apparently extinct and doubtfully lichenized fungus, typically classified in thePyrenulaceae. Anatomical studies reveal that it is facultatively lichenized. The structure of its hamathecium suggests affinities with Dothideomycetes rather than Eurotiomycetes. Molecular analysis using nuLSU and mtSSU markers demonstrates for the first time its inclusion inTrypetheliaceae, outside the core genera as part of the early diverging lineages in this family. The known distribution ofPolypyrenulais extended to Mexico and South America, new information on its phorophyte associations is provided, and the namePolypyrenula sexlocularisis reinstated as the correct name for this species.

Ocellularia (Graphidaceae) is a genus of crustose lichens comprising c.200 species, four known from Australia and New Zealand. Based on recent collections from northern North Island, we describe a ...

The ‘competition-relatedness’ hypothesis postulates that co-occurring taxa should be more distantly related, because of lower competition. This hypothesis has been criticized for its dependence on untested assumptions and its exclusion of other assembly forces beyond competition and habitat filtering to explain the co-existence of closely related taxa. Here we analyzed the patterns of co-occurring individuals of lichenized fungi in theGraphis scriptacomplex, a monophyletic group of species occurring in temperate forests throughout the Northern Hemisphere. We generated sequences for three nuclear ribosomal and protein markers (nuLSU,RPB2,EF-1) and combined them with previously generated sequences to reconstruct an updated phylogeny for the complex. The resulting phylogeny was used to determine the patterns of co-occurrences at regional and at sample (tree) scales by calculating standard effect size of mean pairwise distance (SES.MPD) among co-occurring samples to determine whether they were more clustered than expected from chance. The resulting phylogeny revealed multiple distinct lineages, suggesting the presence of several phylogenetic species in this complex. At the regional and local (site) levels, SES.MPD exhibited significant clustering for five out of six regions. The sample (tree) scale SES. MPD values also suggested some clustering but the corresponding metrics did not deviate significantly from the null expectation. The differences in the SES.MPD values and their significance indicated that habitat filtering and/or local diversification may be operating at the regional level, while the local assemblies on each tree are interpreted as being the result of local competition or random colonization.

Lichens are well known as pioneer organisms colonizing bare surfaces such as rocks and therefore have been hypothesized to play a role in the early formation of terrestrial ecosystems. Given the rarity of fossil evidence, our understanding of the evolutionary history of lichen-forming fungi is primarily based on molecular dating approaches. These studies suggest extant clades of macrolichens diversified after the K–Pg boundary. Here we corroborate the mid-Mesozoic fossil Daohugouthallus ciliiferus as an epiphytic macrolichen that predates the K-Pg boundary by 100 Mys. Based on new material and geometric morphometric analysis, we demonstrate that the Jurassic fossil is morphologically most similar to Parmeliaceae, but cannot be placed in Parmeliaceae or other similar family-level clades forming macrolichens as these evolved much later. Consequently, a new family, Daohugouthallaceae, is proposed here to accommodate this fossil, which reveals macrolichens may have been diverse long before the Cenozoic diversification of extant lineages.

Subandine and high Andean forests are often dominated by oak trees and these are recognized as suitable phorophytes for diverse epiphyte communities. Among the latter, lichens in the upper strata appear to be understudied. Here, we report the discovery of two new species of Astrothelium (Trypetheliaceae), a diverse genus of tropical crustose lichens, both with the unusual feature of muriform, amyloid ascospores. Astrothelium mordonialensis is characterized by an olive-green thallus with solitary ascomata producing muriform and amyloid ascospores with tightened center and A. rogitamae by a light brownish to greenish grey thallus with yellow pigmented solitary ascomata, producing muriform and amyloid ascospores.

The early-successional status of lichens in modern terrestrial ecosystems, together with the role lichen-mediated weathering plays in the carbon cycle, have contributed to the long and widely held assumption that lichens occupied early terrestrial ecosystems prior to the evolution of vascular plants and drove global change during this time. Their poor preservation potential and the classification of ambiguous fossils as lichens or other fungal-algal associations have further reinforced this view. As unambiguous fossil data are lacking to demonstrate the presence of lichens prior to vascular plants, we utilize an alternate approach to assess their historic presence in early terrestrial ecosystems. Here, we analyze new time-calibrated phylogenies of ascomycete fungi and chlorophytan algae, that intensively sample lineages with lichen symbionts. Age estimates for several interacting clades show broad congruence and demonstrate that fungal origins of lichenization postdate the earliest tracheophytes. Coupled with the absence of unambiguous fossil data, our work finds no support for lichens having mediated global change during the Neoproterozoic-early Paleozoic prior to vascular plants. We conclude by discussing our findings in the context of Neoproterozoic-Paleozoic terrestrial ecosystem evolution and the paleoecological context in which vascular plants evolved.

We provide an updated survey for Sri Lanka of species of Graphis sensu Staiger, recently divided into Graphis s. str. and Allographa, including brief descriptions and a key to all 124 species currently known. Six new species are described: Allographa bambusicola Weerakoon, Lücking & Aptroot, a bambusicolous Allographa with entire labia, a laterally carbonized excipulum, 80–100 × 15–17 µm large, muriform ascospores and a rather thick, irregularly verrucose lateral thalline margin of the lirellae; A. weerasooriyana Weerakoon, Arachchige & Lücking, a corticolous Allographa resembling A. rustica Kremp. in overall anatomy and chemistry, but with a verrucose thalline margin of the lirellae and labia not distinctly raised above the thalline margin; Graphis flosculifera Weerakoon, Lücking & Aptroot, a corticolous Graphis resembling G. insulana but differing in the unique disposition of the lirellae and the slightly more elongate ascospores; G. rajapakshana Weerakoon, Lücking & Aptroot, a corticolous Graphis resembling G. desquamescens, including in ascospore size, but with lirellae with a distinct lateral thalline margin; G. rimosothallina Weerakoon, Lücking & Aptroot, a corticolous Graphis with a thick, uneven, rimose thallus and Fissurina-like lirellae, a completely carbonized excipulum and transversely 7-septate ascospores, 32–37 × 8–10 µm; and G. thunsinhalayensis Weerakoon, Arachchige & Lücking, a corticolous Graphis resembling G. subalbostriata but with smaller ascospores and lacking white lines between the striae of the labia. We also validate the name G. verrucoserpens Lücking. A total of 106 species are reported here for the first time from Sri Lanka. A biogeographical comparison with two other well-sampled countries (Costa Rica and Thailand) revealed a significantly higher similarity in species composition with Costa Rica than between Thailand and Costa Rica, suggesting a potential signature of the ‘biotic ferry’ hypothesis, that is the migration of lineages from Gondwana (partly corresponding to the modern Neotropics) via the north-eastwards drifting Indian subcontinent and subsequent interchange with Laurasia (partly corresponding to the modern eastern Paleotropics). However, the evolutionary timeline of the clades involved does not support this hypothesis and suggests an alternative explanation of geologically more recent mid- to long-distance dispersal.

Classification is the most important approach to cataloging biological diversity. It serves as a principal means of communication between scientific disciplines, as well as between scientis...

The genus Graphis sensu Staiger was recently divided into two genera, Graphis s. str. and Allographa. The latter contains mostly species with robust lirellae with a well-developed, often massively carbonized excipulum. With one exception, it also contains all species with a pigmented, yellow to orange pruina on the lirellae. Until now, seven species of Allographa were known with this character, all present in the Neotropics and one also in Africa. Here we describe two further species, both from tropical Asia, thus extending the known distribution of Allographa species with pigmented lirellae to the entire tropics. Allographa kamojangensis Jatnika, Noer & Lücking sp. nov. from Indonesia (Java) was recognized as a new taxon on the social media Facebook site Lichens Connecting People. Detailed studies showed that it deviates from the neotropical A. firferi in the much larger ascospores and the orange, K+ immediately purple-violet pigment, and from A. lutea in the completely carbonized excipulum and the larger ascospores. Allographa jayatilakana Weerakoon, Arachchige & Lücking sp. nov. was discovered in the second author's backyard during a recent inventory of Graphidaceae in Sri Lanka. It differs from A. flavominiata in the much shorter ascospores, from A. firferi in the terminally muriform ascospores, and from A. ochracea in the yellow-orange, K+ yellow then slowly purple-violet pruina. A key is presented to all nine species of Allographa with pigmented lirellae.

Presentamos un primer estudio de la diversidad y composicion de comunidades de liquenes en remanentes de bosque seco tropical (bs-T) en el departamento de Atlantico, Colombia. Se muestrearon liquenes en dos de las tres areas protegidas del departamento: Distrito de Manejo Integrado (DMI) Luriza y Reserva Forestal Protectora (RFP) El Palomar. El inventario registro 61 especies, incluyendo cuatro especies nuevas para la ciencia: Fissurina linoana Lucking, Moncada & G. Rodr. sp. nov., Graphis lurizana Lucking, Moncada & Celis sp. nov., Graphis mokanarum Lucking, Moncada & M.C. Martinez sp. nov. y Phaeographis galeanoae Lucking, Moncada & B. Salgado-N. sp. nov. Arthonia erupta y Coenogonium saepincola son nuevos registros para America del Sur, mientras que trece especies son reportadas por primera vez para Colombia. Unas 37 especies adicionales son nuevos reportes para Atlantico, incrementando el total de especies conocidas para el departamento de 27 a 84. Con 42 especies en Luriza y 31 en El Palomar, la riqueza de especies fue comparable con la de otros sitios de bs-T en el Neotropico. El traslape en la composicion de especies entre los dos sitios es bajo, con solamente doce (20 %) compartidas, indicando un alto nivel de heterogeneidad. Las afinidades biogeograficas son mayores con los bs-T de Centroamerica, lo que coincide con las de plantas lenosas. Estos resultados enfatizan la importancia de los remanentes de bs-T en Colombia para la conservacion de una biodiversidad poco conocida.

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.

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.

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.

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.

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.

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.

In the framework of continuing studies on the Graphidaceae in China, the status of all taxa traditionally assigned to the genus Graphina reported from China are resolved in the present paper. Five new combinations are made, namely Diorygma isabellinum (Zahlbr.) Z.F. Jia & Lücking, comb. nov., Fissurina adscribens (Nyl.) Z.F. Jia & Lücking, comb. nov., Graphis lecanactiformis (Zahlbr.) Z.F. Jia & Lücking, comb. nov., Phaeographis haloniata (Zahlbr.) Z.F. Jia & Lücking, comb. nov. and Platygramme taiwanensis (J.C. Wei) Z.F. Jia & Lucking, comb. nov. Five new synonymies were found: Graphina olivascens Zahlbr. (= Fissurina adscribens), Graphina plumbicolor Zahlbr. (= Phaeographis haloniata), Graphina roridula Zahlbr. and its variety platypoda Zahlbr. [= Diorygma pachygraphum (Nyl.) Kalb, Staiger & Elix], and Graphina taiwanensis f. obscurata J.C. Wei (= Platygramme taiwanensis).

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.

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.

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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 revisionary synopsis is presented for the familyTrypetheliaceae, based on a separately published phylogenetic analysis of a large number of species, morpho-anatomical and chemical study of extensive material, and revision of numerous type specimens. A total of 418 species is formally accepted in this synopsis, distributed among 15 genera as follows:Aptrootia(3),Architrypethelium(7),Astrothelium(242),Bathelium(16),Bogoriella(29),Constrictolumina(9),Dictyomeridium(7),Distothelia(3),Marcelaria(3),Nigrovothelium(2),Novomicrothelia(1),Polymeridium(50),Pseudopyrenula(20),Trypethelium(16), andViridothelium(10). All accepted genera, including new genera described separately in this issue, are keyed out and briefly described and discussed, and keys are provided for all accepted species within each genus. Entries with full synonymy and brief descriptions, and in part also discussions, are provided for all accepted species, except those newly described elsewhere in this issue, which are cross-referenced in the corresponding keys. The description of the newly defined genera takes into account phylogeny in combination with morpho-anatomical features with the result that they are mostly recognizable by a combination of thallus, ascoma and ascospore features. Most species previously assigned to the generaAstrothelium,Campylothelium,Cryptothelium, andTrypethelium, based on a schematic concept of ascoma morphology and ascospore septation, are now included in a single genus,Astrothelium, with highly variable ascoma morphology and ascospore septation but invariably with astrothelioid ascospores (at least when young), that is diamond-shaped lumina, and a well-developed, corticate, usually olive-green thallus that often covers the ascomata. While the generaAptrootia(large, brown, muriform ascospores),Architrypethelium(large, mostly 3-septate ascospores), andPseudopyrenula(ecorticate, white thalli and astrothelioid ascospores) are maintained,Trypetheliumis redefined to include species with raised, pseudostromatic ascomata and multiseptate ascospores with thin septa. The sister group ofTrypetheliumis the genusMarcelaria, with brightly coloured pseudostromata and muriform ascospores.Batheliumis now limited to species with strongly raised, fully exposed pseudostromata and septate to muriform ascospores with thin septa. Several genera are recognized for more basal lineages with mostly ecorticate, white thalli and solitary, exposed ascomata previously assigned toArthopyrenia,MycomicrotheliaandPolymeridium, viz. Bogoriella,Constrictolumina,Dictyomeridium, andNovomicrothelia. In addition, separate genera are accepted for theTrypethelium tropicum(Nigrovothelium) andT. virens(Viridothelium) groups. In addition, a refined species concept resulting from phylogenetic studies is employed which pays particular attention to morphological features of the thallus and ascomata. Of a total of 526 names checked, 107 remain synonyms of accepted names and a further eight are newly excluded from the family. Based on these redispositions, the following 146 new combinations are proposed, including reinstatement of numerous names previously subsumed into synonymy:Architrypethelium columbianum(Nyl.) Aptroot & Lücking comb. nov.,A. grande(Kremp.) Aptroot & Lücking comb. nov.,Astrothelium aeneum(Eschw.) Aptroot & Lücking comb. nov.,A. alboverrucum(Makhija & Patw.) Aptroot & Lücking comb. nov.,A. amazonum(R. C. Harris) Aptroot & Lücking comb. nov.,A. ambiguum(Malme) Aptroot & Lücking comb. nov.,A. andamanicum(Makhija & Patw.) Aptroot comb. nov.,A. annulare(Spreng.) Aptroot & Lücking comb. nov.,A. aurantiacum(Makhija & Patw) Aptroot & Lücking comb. nov.,A. auratum(R. C. Harris) Aptroot & Lücking comb. nov.,A. aureomaculatum(Vain.) Aptroot & Lücking comb. nov.,A. basilicum(Kremp.) Aptroot & Lücking comb. nov.,A. bicolor(Taylor) Aptroot & Lücking comb. nov.,A. buckii(R. C. Harris) Aptroot & Lücking comb. nov.,A. calosporum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. cartilagineum(Fée) Aptroot & Lücking comb. nov.,A. cecidiogenum(Aptroot & Lücking) Aptroot & Lücking comb. nov.,A. ceratinum(Fée) Aptroot & Lücking comb. nov.,A. chapadense(Malme) Aptroot & Lücking comb. nov.,A. chrysoglyphum(Vain.) Aptroot & Lücking comb. nov.,A. chrysostomum(Vain.) Aptroot & Lücking comb. nov.,A. cinereorosellum(Kremp.) Aptroot & Lücking comb. nov.,A. cinereum(Müll. Arg.) Aptroot & Lücking comb. et stat. nov.,A. confluens(Müll. Arg.) Aptroot & Lücking comb. nov.,A. consimile(Müll. Arg.) Aptroot & Lücking comb. nov.,A. deforme(Fée) Aptroot & Lücking comb. nov.,A. defossum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. degenerans(Vain.) Aptroot & Lücking comb. nov.,A. dissimilum(Makhija & Patw.) Aptroot & Lücking comb. nov.,A. effusum(Aptroot & Sipman) Aptroot & Lücking comb. nov.,A. endochryseum(Vain.) Aptroot & Lücking comb. nov.,A. exostemmatis(Müll. Arg.) Aptroot & Lücking comb. nov.,A. feei(C. F. W. Meissn.) Aptroot & Lücking comb. nov.,A. ferrugineum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. galligenum(Aptroot) Aptroot & Lücking comb. nov.,A. gigantosporum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. indicum(Upreti & Ajay Singh) Aptroot & Lücking comb. nov.,A. infossum(Nyl.) Aptroot & Lücking comb. nov.,A. infuscatulum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. irregulare(Müll. Arg.) Aptroot & Lücking comb. nov.,A. keralense(Upreti & Ajay Singh) Aptroot & Lücking comb. nov.,A. kunzei(Fée) Aptroot & Lücking comb. nov.,A. leioplacum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. lugescens(Nyl.) Aptroot & Lücking comb. nov.,A. luridum(Zahlbr.) Aptroot & Lücking comb. nov.,A. macrocarpum(Fée) Aptroot & Lücking comb. nov.,A. macrosporum(Makhija & Patw.) Aptroot & Lücking comb. nov.,A. marcidum(Fée) Aptroot & Lücking comb. nov.,A. megaleium(Kremp.) Aptroot & Lücking comb. nov.,A. megalophthalmum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. megalostomum(Vain.) Aptroot & Lücking comb. nov.,A. megaspermum(Mont.) Aptroot & Lücking comb. nov.,A. meiophorum(Nyl.) Aptroot & Lücking comb. nov.,A. meristosporoides(P. M. McCarthy & Vongshew.) Aptroot & Lücking comb. nov.,A. meristosporum(Mont. & Bosch) Aptroot & Lücking comb. nov.,A. neogalbineum(R. C. Harris) Aptroot & Lücking comb. nov.,A. nigratum(Müll. Arg.) Aptroot & Lücking comb. et stat. nov.,A. nigrorufum(Makhija & Patw.) Aptroot & Lücking comb. nov.,A. nitidiusculum(Nyl.) Aptroot & Lücking comb. nov.,A. octosporum(Vain.) Aptroot & Lücking comb. nov.,A. oligocarpum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. olivaceofuscum(Zenker) Aptroot & Lücking comb. nov.,A. papillosum(P. M. McCarthy) Aptroot & Lücking comb. nov.,A. papulosum(Nyl.) Aptroot & Lücking comb. nov.,A. peranceps(Kremp.) Aptroot & Lücking comb. nov.,A. phaeothelium(Nyl.) Aptroot & Lücking comb. nov.,A. phlyctaenua(Fée) Aptroot & Lücking comb. nov.,A. porosum(Ach.) Aptroot & Lücking comb. nov.,A. praetervisum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. pseudoplatystomum(Makhija & Patw.) Aptroot & Lücking comb. nov.,A. pseudovariatum(Upreti & Ajay Singh) Aptroot & Lücking comb. nov.,A. puiggarii(Müll. Arg.) Aptroot & Lücking comb. nov.,A. pulcherrimum(Fée) Aptroot & Lücking comb. nov.,A. pupula(Ach.) Aptroot & Lücking comb. nov.,A. purpurascens(Müll. Arg.) Aptroot & Lücking comb. nov.,A. pustulatum(Vain.) Aptroot & Lücking comb. nov.,A. rufescens(Müll. Arg.) Aptroot & Lücking comb. et stat. nov.,A. sanguinarium(Malme) Aptroot & Lücking comb. nov.,A. santessonii(Letr.-Gal.) Aptroot & Lücking comb. nov.,A. saxicola(Malme) Aptroot & Lücking comb. nov.,A. scoria(Fée) Aptroot & Lücking comb. nov.,A. scorizum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. sierraleonense(C. W. Dodge) Aptroot & Lücking comb. nov.,A. sikkimense(Makhija & Patw.) Aptroot & Lücking comb. nov.,A. spectabile(Aptroot & Ferraro) Aptroot & Lücking comb. nov.,A. sphaerioides(Mont.) Aptroot & Lücking comb. nov.,A. stramineum(Malme) Aptroot & Lücking comb. nov.,A. straminicolor(Nyl.) Aptroot & Lücking comb. nov.,A. subcatervarium(Malme) Aptroot & Lücking comb. nov.,A. subdiscretum(Nyl.) Aptroot & Lücking comb. nov.,A. subdisjunctum(Müll. Arg.) Aptroot & Lücking comb. nov.,A. subdissocians(Nyl. ex Vain.) Aptroot & Lücking comb. et stat. nov.,A. superbum(Fr.) Aptroot & Lücking comb. nov.,A. tenue(Aptroot) Aptroot & Lücking comb. nov.,A. thelotremoides(Nyl.) Aptroot & Lücking comb. nov.,A. trypethelizans(Nyl.) Aptroot & Lücking comb. nov.,A. tuberculosum(Vain.) Aptroot & Lücking comb. nov.,A. ubianense(Vain.) Aptroot & Lücking comb. nov.,A. variatum(Nyl.) Aptroot & Lücking comb. nov.,A. vezdae(Makhija & Patw.) Aptroot & Lücking comb. nov.,Bathelium austroafricanum(Zahlbr.) Aptroot & Lücking comb. nov.,B. nigroporum(Makhija & Patw.) Aptroot & Lücking comb. nov.,Bogoriella alata(Groenh. ex Aptroot) Aptroot & Lücking comb. nov.,B. annonacea(Müll. Arg.) Aptroot & Lücking comb. nov.,B. apposita(Nyl.) Aptroot & Lücking comb. nov.,B. captiosa(Kremp.) Aptroot & Lücking comb. nov.,B. collospora(Vain.) Aptroot & Lücking comb. nov.,B. confluens(Müll. Arg.) Aptroot & Lücking comb. nov.,B. conothelena(Nyl.) Aptroot & Lücking comb. nov.,B. decipiens(Müll. Arg.) Aptroot & Lücking comb. nov.,B. exigua(Müll. Arg.) Aptroot & Lücking comb. nov.,B. fumosula(Zahlbr.) Aptroot & Lücking comb. nov.,B. hemisphaerica(Müll. Arg.) Aptroot & Lücking comb. nov.,B. lateralis(Sipman) Aptroot & Lücking comb. nov.,B. leuckertii(D. Hawksw. & J. C. David) Aptroot & Lücking comb. nov.,B. macrocarpa(Komposch, Aptroot & Hafellner) Aptroot & Lücking comb. nov.,B. megaspora(Aptroot & M. Cáceres) Aptroot & Lücking comb. nov.,B. miculiformis(Nyl. ex Müll. Arg.) Aptroot & Lücking comb. nov.,B. minutula(Zahlbr.) Aptroot & Lücking comb. nov.,B. modesta(Müll. Arg.) Aptroot & Lücking comb. nov.,B. nonensis(Stirt.) Aptroot & Lücking comb. nov.,B. obovata(Stirt.) Aptroot & Lücking comb. nov.,B. pachytheca(Sacc. & Syd.) Aptroot & Lücking comb. nov.,B. punctata(Aptroot) Aptroot & Lücking comb. nov.,B. queenslandica(Müll. Arg.) Aptroot & Lücking comb. nov.,B. socialis(Zahlbr.) Aptroot & Lücking comb. nov.,B. striguloides(Sérus. & Aptroot) Aptroot & Lücking comb. nov.,B. subfallens(Müll. Arg.) Aptroot & Lücking comb. nov.,B. thelena(Ach.) Aptroot & Lücking comb. nov.,B. triangularis(Aptroot) Aptroot & Lücking comb. nov.,B. xanthonica(Komposch, Aptroot & Hafellner) Aptroot & Lücking comb. nov.,Constrictolumina esenbeckiana(Fée) Lücking, M. P. Nelsen & Aptroot comb. nov.,C. leucostoma(Müll. Arg.) Lücking, M. P. Nelsen & Aptroot comb. nov.,C. lyrata(R. C. Harris) Lücking, M. P. Nelsen & Aptroot comb. nov.,C. majuscula(Nyl.) Lücking, M. P. Nelsen & Aptroot comb. nov.,C. malaccitula(Nyl.) Lücking, M. P. Nelsen & Aptroot comb. nov.,C. porospora(Vain.) Lücking, M. P. Nelsen & Aptroot comb. nov.,Dictyomeridium amylosporum(Vain.) Aptroot, M. P. Nelsen & Lücking comb. nov.,D. campylothelioides(Aptroot & Sipman) Aptroot, M. P. Nelsen & Lücking comb. nov.,D. immersum(Aptroot, A. A. Menezes & M. Cáceres) Aptroot, M. P. Nelsen & Lücking comb. nov.,D. isohypocrellinum(Xavier-Leite, M. Cáceres & Aptroot) Aptroot, M. P. Nelsen & Lücking comb. nov.,D. paraproponens(Aptroot, M. Cáceres & E. L. Lima) Aptroot, M. P. Nelsen & Lücking comb. nov.,Distothelia rubrostoma(Aptroot) Aptroot & Lücking comb. nov.,Phyllobathelium chlorogastricum(Müll. Arg.) Aptroot & Lücking comb. nov.,Pseudopyrenula cubana(Müll. Arg.) Aptroot & Lücking comb. nov.,Viridothelium cinereoglaucescens(Vain.) Lücking, M. P. Nelsen & Aptroot comb. nov.,V. indutum(Stirt.) Aptroot & Lücking comb. nov., andV. megaspermum(Makhija & Patw.) Aptroot & Lücking comb. nov. In addition, six replacement names are proposed:Astrothelium campylocartilagineumAptroot & Lücking nom. nov.,A. grossoidesAptroot & Lücking nom. nov.,A. octosporoidesAptroot & Lücking nom. nov.,A. scoriotheliumAptroot & Lücking nom. nov.,A. pyrenastrosulphureumAptroot & Lücking nom. nov., andBathelium pruinolucensAptroot & Lücking nom. et stat. nov. Along with this, 57 lectotypes are newly designated. Most species (392 out of 418) are illustrated, with a total of 697 images in 59 plates, including 406 type specimens. Where appropriate, taxa are briefly discussed. New country or continental records are listed for many species in their revised circumscription. A checklist of taxa described or placed in genera belonging inTrypetheliaceaebut previously excluded from the family, and their current names, is also provided.

Following a large-scale phylogenetic study of the lichenized genus Cora (Basidiomycota: Agaricales: Hygrophoraceae), we formally describe 70 new species, honouring the seventieth birthday of David Leslie Hawksworth, one of the preeminent figures in mycology and lichenology in the past 50 years. Based on an updated phylogeny using the ITS fungal barcoding locus, we now recognize 189 taxa in a genus that until recently was considered to represent a single species; including this contribution, 92 of these are formally recognized, including five taxa based on historical names or collections that have not been sequenced. Species of Cora can be recognized by a combination of morphological (size, colour, lobe configuration, surface hairs, hymenophore size and shape), anatomical (thallus thickness, cortex structure, photobiont type, hyphal papillae), and ecogeographical features (substrate, habitat, distribution), and a keytable allowing the identification of all accepted taxa is provided. The new species are: Cora accipiter Moncada, Madrinan & Lucking spec. nov., C. applanata Moncada, Soto-Medina & Lucking spec. nov., C. arachnodavidea Moncada, Dal Forno & Lucking spec. nov., C. arborescens Dal Forno, Chaves & Lucking spec. nov., C. arcabucana Moncada, C. Rodriguez & Lucking spec. nov., C. aturucoa Lucking, Moncada & C. Vargas spec. nov., C. auriculeslia Moncada, Yanez-Ayabaca & Lucking spec. nov., C. barbifera Moncada, Patino & Lucking spec. nov., C. boleslia Lucking, E. Morales & Dal Forno spec. nov., C. caliginosa Holgado, Rivas Plata & Perlmutter spec. nov., C. campestris Dal Forno, Eliasaro & Spielmann spec. nov., C. canari Nugra, Dal Forno & Lucking spec. nov., C. caraana Lucking, Martins & Lucheta spec. nov., C. casasolana Moncada, R.-E. Perez & Lucking spec. nov., C. caucensis Moncada, M. Gut. & Lucking spec. nov., C. celestinoa Moncada, Cabrera-Amaya & Lucking spec. nov., C. comaltepeca Moncada, R.-E. Perez & Herrera-Camp. spec. nov., C. corani Lucking, E. Morales & Dal Forno spec. nov., C. corelleslia Moncada, A. Suarez-Corredor & Lucking spec. nov., C. crispoleslia Moncada, J. Molina & Lucking spec. nov., C. cuzcoensis Holgado, Rivas Plata & Perlmutter spec. nov., C. dalehana Moncada, Madrinan & Lucking spec. nov., C. davibogotana Lucking, Moncada & Coca spec. nov., C. davicrinita Moncada, Madrinan & Lucking spec. nov., C. davidia Moncada, L. Vargas & Lucking spec. nov., C. dewisanti Moncada, A. Suarez-Corredor & Lucking spec. nov., C. dulcis Moncada, R.-E. Perez & Lucking spec. nov., C. elephas Lucking, Moncada & L. Vargas spec. nov., C. fuscodavidiana Lucking, Moncada & L. Vargas spec. nov., C. garagoa Simijaca, Moncada & Lucking spec. nov., C. gigantea Lucking, Moncada & Coca spec. nov., C. gomeziana Dal Forno, Chaves & Lucking spec. nov., C. guajalitensis Lucking, Robayo & Dal Forno spec. nov., C. hafecesweorthensis Moncada, Lucking & R. Pelaez spec. nov., C. haledana Dal Forno, Chaves & Lucking spec. nov., C. hawksworthiana Dal Forno, P. Nelson & Lucking spec. nov., C. hochesuordensis Lucking, E. Morales & Dal Forno spec. nov., C. hymenocarpa Lucking, Chaves & Lawrey spec. nov., C. imi Lucking, Chaves & Lawrey spec. nov., C. itabaiana Dal Forno, Aptroot & M. Caceres spec. nov., C. leslactuca Lucking, Moncada & R. Pelaez spec. nov., C. maxima Wilk, Dal Forno & Lucking spec. nov., C. minutula Lucking, Moncada & Yanez-Ayabaca spec. nov., C. palaeotropica Weerakoon, Aptroot & Lucking spec. nov., C. palustris Dal Forno, Chaves & Lucking spec. nov., C. parabovei Dal Forno, Kukwa & Lucking spec. nov., C. paraciferrii Lucking, Moncada & J.E. Hern. spec. nov., C. paraminor Dal Forno, Chaves & Lucking spec. nov., C. pastorum Moncada, Patino & Lucking spec. nov., C. pichinchensis Paredes, Jonitz & Dal Forno spec. nov., C. pikynasa J.-M. Torres, Moncada & Lucking spec. nov., C. pseudobovei Wilk, Dal Forno & Lucking spec. nov., C. pseudocorani Lucking, E. Morales & Dal Forno spec. nov., C. putumayensis L.J. Arias, Moncada & Lucking spec. nov., C. quillacinga Moncada, F. Ortega & Lucking spec. nov., C. rothesiorum Moncada, Madrinan & Lucking spec. nov., C. rubrosanguinea Nugra, Moncada & Lucking spec. nov., C. santacruzensis Dal Forno, Bungartz & Yanez-Ayabaca, spec. nov., C. schizophylloides Moncada, C. Rodriguez & Lucking spec. nov., C. smaragdina Lucking, Rivas Plata & Chaves spec. nov., C. soredavidia Dal Forno, Marcelli & Lucking spec. nov., C. subdavicrinita Moncada, J. Molina & Lucking spec. nov., C. suturifera Nugra, Besal & Lucking spec. nov., C. terrestris Dal Forno, Chaves & Lucking spec. nov., C. terricoleslia Wilk, Dal Forno & Lucking spec. nov., C. udebeceana Moncada, R. Pelaez & Lucking spec. nov., C. urceolata Moncada, Coca & Lucking spec. nov., C. verjonensis Lucking, Moncada & Dal Forno spec. nov., C. viliewoa Lucking, Chaves & Soto-Medina spec. nov., and C. yukiboa Mercado-Diaz, Moncada & Lucking spec. nov. Furthermore, the taxonomic status of the recently described or recognized species C. arachnoidea, C. aspera, C. ciferrii, and C. reticulifera, is revised.

The new name Neosergipea M. Cáceres, Ertz & Aptroot is introduced to replace Sergipea M. Cáceres, Ertz & Aptroot, which is a later homonym of Sergipea Regali, Uesugui & Santos, a genus of fossil pollen. Using the small subunit of the mitochondrial rDNA cistron, we present an updated phylogeny of the Enterographa clade in Roccellaceae which includes the genera Dichosporidium, Enterographa, Erythrodecton, Mazosia, and Neosergipea. While in a previous analysis the relationship between Neosergipea and Dichosporidium was unresolved, our results suggest Neosergipea to be an unsupported sister to Dichosporidium s. lat. The latter potentially represents two distinct genera, differing in ascospore type.

The aim of this study was to evaluate the spatial effect of a natural source of sulphur pollution on the species diversity, richness and distribution of corticolous lichens in a páramo zone at the mine ‘El Vinagre’ (Puracé, Cauca, Colombia). Three zones at different distances from the pollution source were established: zone 1 with a high degree of contamination, a potentially mildly affected or transitional zone 2, and a zone 3 free of disturbance. In each zone, 10 phorophytes ofWeinmannia microphylla(Cunoniaceae) were sampled, and all lichens in a 150 cm vertical transect 50 cm above the ground were collected and identified. Phorophyte parameters were measured (bark pH and diameter at breast height) and the samples were georeferenced. In order to evaluate the impact on lichens, non-metric multidimensional scaling (NMS) and indicator species analysis were carried out. ANOVA and Spearman correlations were performed to assess the relationships of environmental and tree variables between zones and with lichen community structure. The index of atmospheric purity (IAP) and the environmental rating factor (FCA) were evaluated for the three zones. In total, 104 lichen species were recorded, of which 72 were identified to species, 17 to genus, and four to family; 11 samples could not be identified. NMS clustered samples according to zone and the main axis which were correlated with bark pH and distance from pollution source. We found eight indicator species characterizing different zones, and four marginally significant indicator species. Using the IAP, we established three isocontamination areas, with zones 2 and 3 classified as more or less pristine zones and zone 1 as a polluted zone (supported by bark pH as a proxy for current pollution). Diversity was lowest in zone 1, closest to the pollution source, and lichen species composition differed between zones. Differences between lichens in zones 2 and 3 appear mostly unrelated to the current pollution source and might be more related to historical differences in impact from a 20-year-old pollution source.

Species in the fungal genus Sticta form symbiotic associations primarily with either green algae or cyanobacteria, but tripartite associations or photosymbiodemes involving both types of photobionts occur in some species. Sticta is known to associate with green algae in the genus Symbiochloris. However, previous studies have shown that algae from other genera, such as Heveochlorella, may also be suitable partners for Sticta. We examined the diversity of green algal partners in the genus Sticta and assessed the patterns of association between the host fungus and its algal symbiont. We used multi-locus sequence data from multiple individuals collected in Australia, Cuba, Madagascar, Mauritius, New Zealand, Reunion and South America to infer phylogenies for fungal and algal partners and performed tests of congruence to assess coevolution between the partners. In addition, event-based methods were implemented to examine which cophylogenetic processes have led to the observed association patterns in Sticta and its green algal symbionts. Our results show that in addition to Symbiochloris, Sticta associates with green algae from the genera Chloroidium, Coccomyxa, Elliptochloris and Heveochlorella, the latter being the most common algal symbiont associated with Sticta in this study. Geography plays a strong role in shaping fungal-algal association patterns in Sticta as mycobionts associate with different algal lineages in different geographic locations. While fungal and algal phylogenies were mostly congruent, event-based methods did not find any evidence for cospeciation between the partners. Instead, the association patterns observed in Sticta and associated algae, were largely explained by other cophylogenetic events such as host-switches, losses of symbiont and failure of the symbiont to diverge with its host. Our results also show that tripartite associations with green algae evolved multiple times in Sticta.

During ecological studies on corticolous lichens in Restinga vegetation in northeastern Brazil, many species of crustose Caliciaceae formerly placed in the collective genus Buellia s.l. were collected. A new species was discovered and is described here—Gassicurtia restingiana D.S.Andrade, M.Caceres & Aptroot differs from G. catasema in the absence of lichexanthone. Two identification keys are provided, one for all species of Gassicurtia known from Brazil, and one for crustose, epiphytic, non-mazaediate species of Caliciaceae found in Restinga areas in northeastern Brazil.

We report the production of the perylenequinone pigment elsinochrome A in aposymbiotic culture of the mycobiont of the crustose epiphytic lichen Graphis elongata Zenker (Lecanoromycetes), collected in Argentina (Buenos Aires). The substance was not detected in the lichenized thallus (using HPLC techniques) and is otherwise only known from one unrelated lichen and a few genera of non-lichenized, plant-pathogenic fungi in the class Dothideomycetes. The phylogenetic affinities of the lichen mycobiont and the cultured fungus were confirmed using DNA sequence data of the mitochondrial small subunit rDNA (mtSSU), which place the lichen fungus into the Allographa clade within Graphidaceae. The mycobiont pigment was purified and characterized by spectroscopic methods. This is the first case where a rare pigment, otherwise known from non-lichenized, plant-pathogenic fungi, is produced in aposymbiotic culture of a lichen mycobiont, while, at the same time, being absent from the lichen thallus itself. Based on this finding, we discuss the previously postulated hypothesis that lichen mycobionts maintain secondary metabolic pathways of non-lichenized ancestors in their genome, while gene expression and production of metabolites is suppressed in the lichenized state due to toxicity to the photobiont. Fil: Fazio, Alejandra Teresa. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Micología y Botánica. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Micología y Botánica; Argentina Fil: Adler, Monica Teresa. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Micología y Botánica. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Micología y Botánica; Argentina Fil: Parnmen, Sittiporn. National Institute of Health. Toxicology Center; Tailandia. Associate Researcher at the Field Museum; Estados Unidos Fil: Lücking, Robert. Associate Researcher at the Field Museum; Estados Unidos. Botanic Garden and Botanical Museum; Alemania Fil: Maier, Marta Silvia. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Unidad de Microanálisis y Métodos Físicos en Química Orgánica. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Unidad de Microanálisis y Métodos Físicos en Química Orgánica; Argentina

Premise of research. With more than 2000 described species, Graphidaceae represents one of the largest families of lichenized fungi. The underlying causes for such a high level of diversity have not yet been studied in detail. Two possible drivers of high diversity in this group include its predominantly tropical habitat, a major factor for high species richness in other groups of organisms, and the presence of columella, a unique trait within this family.Methodology. We employed a combination of comparative methods to investigate possible factors that are associated with the hyperdiverse nature of the family, including modeling evolutionary diversity using stepwise Akaike information criterion (MEDUSA) to identify clades with an increased diversification rate and binary state speciation and extinction (BiSSE) analysis to determine whether diversification rates change in association with selected character states.Pivotal results. BiSSE analysis revealed a significantly higher diversification rate in the l...

One new species and six new records of the crustose lichenized genus Graphis are reported from Thailand. Graphis koratensis Pitakpong, Kraichak & Lücking sp. nov. is characterized by lirelline ascocarps with whitish grey or grey-green pruina along the slit, transversely septate ascospores, and the presence of norstictic acid. Phylogenetic analyses with two loci (mtSSU and nuLSU) show the distinct position of this new species within the genus. Six new records for Thailand are reported, including G. cincta (Pers.) Aptroot, G. jejuensis K. H. Moon et al., G. nigrocarpa Adaw. & Makhija, G. renschiana (Müll. Arg.) Stizenb., G. seminuda Müll. Arg., and G. subserpentina Nyl.

Graphis scripta, or script lichen, is a well-known species of crustose lichenized fungi, widely distributed in the temperate region of the Northern Hemisphere. It is now considered to be a species complex, but because of the lack of secondary chemistry and paucity of measurable morphological characters, species delimitation within the complex has been challenging and is thus far based on apothecium and ascospore morphology. In this study, we employed molecular as well as morphological data to assess phylogenetic structure and delimitation of lineages within the G. scripta complex. We generated sequences for four genetic markers (mtSSU, nuLSU, RPB2, and EF-1) and performed phylogenetic analyses. The resulting trees were used to determine the number of distinct lineages by applying a general mixed Yule-coalescent (GMYC) model and species tree estimation through maximum likelihood (STEM). Our analyses suggest between six and seven putative species within the G. scripta complex. However, these did not correspond to the taxa that were recently distinguished based on apothecium morphology and could not be circumscribed with the morphological characters that were traditionally used in the classification of the complex. Any formal taxonomic treatment will require additional sampling and evaluation of additional traits that potentially can characterize these clades.

Three species of lichen-forming fungi in the Gomphillaceae (Ostropales, Ascomycota) are described as new to science: Gyalideopsis sessile, Echinoplaca basalis and Tricharia duotela. The lichens were found on palm leaves in remnants of native vegetation on the campus of Florida Gulf Coast University and at two other conserved sites in southwest Florida.

Based on collections focusing on Graphidaceae made in 2012 at Bukit Timah Nature Reserve (BTNR) in Singapore, seven new species are described: Astrochapsa sipmanii, differing from A. astroidea in the olive-brown thallus and thick and coarse apothecial pruina; Fissurina duplomarginata, differing from F. insidiosa by the double margin of the lirellae and the more greenish, rough thallus; Graphis bukittimaensis, differing from G. phaeospora in the erumpent lirellae with lateral thalline margin and the consistently 1-spored asci; G. singaporensis, differing from G. novopalmicola in the much smaller ascospores and in the immersed, densely branched lirellae with thin lateral thalline margin; Ocellularia subudupiensis, differing from O. udupiensis in the presence of three unknown secondary substances and in the comparatively rough thallus surface; O. gueidaniana, characterized by small, transversely septate, hyaline ascospores, prominent, carbonized but ecolumellate apothecia, and two unknown chemical substances resembling metabolites in the stictic acid chemosyndrome; and O. rivasplatana, differing from O. exigua in the larger, broad-pored apothecia with black-rimmed margin and filled with a black-topped columella. Nineteen species are recognized as new records for Singapore. The substantial diversity of Graphidaceae in this reserve suggests that many more taxa are awaiting discovery, even in such a highly urbanized location.

In many tropical areas, monospecific tree plantations are replacing natural forest. The ecology of these plantations is quite different from that of natural forests, including the diversity and community structure of vascular and cryptogamic epiphytes. Few studies have looked at the ecology of guilds of epiphytes in plantations versus natural forest. Here, we investigated epiphytic, lirellate species of the family Graphidaceae, the largest family of tropical lichen fungi, which are widely distributed and abundant in tropical regions. We compared species richness and community structure in a monospecific plantation of the introduced tree species Eucalyptus globulus versus native oak forest dominated by Quercus humboldtii. Overall species richness was substantially higher in the natural oak forest (41 vs. 14 species, with eight shared between both stands, for a total of 47), whereas species abundance was significantly higher in the gum plantation. While species richness per tree (alpha diversity) was comparable between both stands, average species turnover between trees within each stand (beta diversity) was significantly higher in the natural oak forest, resulting in substantially higher overall species richness (gamma diversity). We conclude that the monospecific gum plantation exhibits both de-diversification (lower overall species richness) and homogenization (more similar communities between trees) of these epiphytic lichen guilds. This is not an effect of phorophyte diversity since in both stands, only a single tree species each was considered. Among the lichens identified, we detected six new to the Neotropics and 29 new records for Colombia.