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Diversity of lichenized Basidiomycota in Costa Rica. (A) Acantholichen pannarioides P. M. Jørg. (lobes to the right 2 mm wide). (B–D) Dictyonema sericeum (Sw.) Berk.; in (B) filamentous-crustose growth form (f. phyllophilum Parmasto) with white prothallus (image section 10 mm wide); in (C) filamentous-pilose growth form [f. schenkianum (M  ̈ll. Arg.) Parmasto] with numerous resupinate basidiocarps (image section 30 mm wide); in (D) filamentous-lobate growth form (f. sericeum ) on a twig (thallus 40 mm wide). (E–F) Dictyonema glabratum (Spreng.) D. Hawksw.; in (E) terricolous, in (F) epiphytic growth form (large lobes 50 mm wide). (G) Cyphellostereum pusiolum (Berk. and M. A. Curtis) D. A. Reid (indistinctly lichenized; basidiocarp 5 mm high). (H) Lepidostroma calocerum (G. W. Martin) Oberw. (basidiocarps 8–10 mm high). (I) Multiclavula sp. (basidiocarps 7–10 mm high). (J–L) M. ichthyiformis ; in (J) thallus with fully hydrated basidiocarps, in (K–L) individual basidiocarps after collecting (basidiocarps 6–9 mm high). All photographs taken in the field by Robert L  ̈cking, except (G) and (K–L) taken in the laboratory. 

Diversity of lichenized Basidiomycota in Costa Rica. (A) Acantholichen pannarioides P. M. Jørg. (lobes to the right 2 mm wide). (B–D) Dictyonema sericeum (Sw.) Berk.; in (B) filamentous-crustose growth form (f. phyllophilum Parmasto) with white prothallus (image section 10 mm wide); in (C) filamentous-pilose growth form [f. schenkianum (M ̈ll. Arg.) Parmasto] with numerous resupinate basidiocarps (image section 30 mm wide); in (D) filamentous-lobate growth form (f. sericeum ) on a twig (thallus 40 mm wide). (E–F) Dictyonema glabratum (Spreng.) D. Hawksw.; in (E) terricolous, in (F) epiphytic growth form (large lobes 50 mm wide). (G) Cyphellostereum pusiolum (Berk. and M. A. Curtis) D. A. Reid (indistinctly lichenized; basidiocarp 5 mm high). (H) Lepidostroma calocerum (G. W. Martin) Oberw. (basidiocarps 8–10 mm high). (I) Multiclavula sp. (basidiocarps 7–10 mm high). (J–L) M. ichthyiformis ; in (J) thallus with fully hydrated basidiocarps, in (K–L) individual basidiocarps after collecting (basidiocarps 6–9 mm high). All photographs taken in the field by Robert L ̈cking, except (G) and (K–L) taken in the laboratory. 

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The new basidiolichen Multiclavula ichthyiformis Nelsen, Lücking, Umaña, Trest & Will-Wolf is described from Costa Rica. The new species differs from other species of Multiclavula in having a basidiocarp with tomentose stipe and flattened lamina with nonamphigenous hymenium. Molecular sequence data (ITS) confirmed its placement within Multiclavula...

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... best-fit model as determined by the Akike information criterion (AIC) in MrModeltest (Nylander, 2004). The analysis was run for 3 000 000 generations with four chains at a ''temperature'' of 0.18, using default settings and sampling every 100 generations. A 50% majority-rule consensus of all sampled post-burn-in trees was constructed in PAUP*. (Fig. 1B-F) Atheliales (Atheliaceae) or euagarics clade Gargas et al., 1995;Hibbett and Thorn, 2001 Largest lichenized genus in Basidiomycota with more than ten species; no recent molecular analysis Cyphellostereum (Fig. 1G Etymology-The name refers to the fishtail-like basidio- carps of the new ...
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... and sampling every 100 generations. A 50% majority-rule consensus of all sampled post-burn-in trees was constructed in PAUP*. (Fig. 1B-F) Atheliales (Atheliaceae) or euagarics clade Gargas et al., 1995;Hibbett and Thorn, 2001 Largest lichenized genus in Basidiomycota with more than ten species; no recent molecular analysis Cyphellostereum (Fig. 1G Etymology-The name refers to the fishtail-like basidio- carps of the new ...
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... terricolous (Fig. 1J), up to 50 mm across but very thin, barely visible as greenish layer, composed of dense colonies of green algae associated with fungal hyphae. Photobiont Coccomyxa ( Fig. 2A-D, G-K), cells broadly ellipsoid, 7-9 3 4-7 lm, each cell individually wrapped by a layer of 1.5-2.0 lm thick fungal hyphae (bulbils), algal cells frequently ...
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... scattered, not aggregate ( Fig. 1J-K), fleshy, unbranched, more or less lanceolate, with a basal stipe 2-3.3-5 mm high and 0.5-0.55-0.7 mm in diameter and an upper widened, fishtail-like lamina 0.6-2.4-4.0 mm high and 0.9- 1.4-1.9 mm wide. Stipe coarsely tomentose, covered by numerous tiny scales (Figs. 1K-L, 2E), pale flesh-colored when moist to translucent white when ...
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... scattered, not aggregate ( Fig. 1J-K), fleshy, unbranched, more or less lanceolate, with a basal stipe 2-3.3-5 mm high and 0.5-0.55-0.7 mm in diameter and an upper widened, fishtail-like lamina 0.6-2.4-4.0 mm high and 0.9- 1.4-1.9 mm wide. Stipe coarsely tomentose, covered by numerous tiny scales (Figs. 1K-L, 2E), pale flesh-colored when moist to translucent white when dry; scales 0.1-0.25-0.4 mm long, tapering, 30-40-50 lm broad at the base and 20-25- 30 lm broad below the apex, formed by densely packed, parallel, sparsely branched hyphae. Both lamina, stipe, and scales frequently producing bulbils enclosing individual algal cells or algal ...

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... P. Kumm [35]. Multiclavula ichthyiformis Nelsen, Lücking, L. Umaña, Trest & Will-Wolf, was identified as a new basidiolichen from Costa Rica with terricolous habits that grows with bryophytes [36]. Psathyrella laurentiana A.H. Sm and Omphalina philonotis (Lasch) Quél. ...
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... Increasing focus was also placed on species recognition in lichenized Basidiomycota, using the fungal ITS barcoding locus (Schoch et al. 2012;Lücking et al. 2014b). As a result, the number of accepted species of lichenized Basidiomycota, including those described herein, has increased dramatically, in particular in the genus Cora ( Fig. 1) (Chaves et al. 2004;Fischer et al. 2007;Nelsen et al. 2007;Hodkinson et al. 2014;Sulzbacher et al. 2012;Yánez et al. 2012;Lücking et al. 2013aLücking et al. , b, 2014bSchmull et al. 2014;Vargas et al. 2014;Yanaga et al. 2015;Dal Forno et al. 2016). This corresponds to an increase of the proportion of Basidiomycota among lichen fungi from approximately 0.2 % until the year 2000 to 0.9 % currently and 1.6 % predicted by the end of 2017. ...
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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...
... Morphologically, Bryoclavula is closely related to Multiclavula R.H. Petersen in having clavarioid basidiocarps and 4-6 sterigmata, and both are lichenised genera. However, Bryoclavula phycophila does not form the globular or bulbil-like lichenised thallus like the species in Multiclavula (Oberwinkler 1970, Nelsen et al. 2007, Masumoto & Degawa 2020a. The two lichenised genera have a distant relationship in the phylogenetic tree according to Masumoto & Degawa (2020a) and also this study. ...
... Notes: Multiclavula is typified by Mu. corynoides and characterised by the simple or branched small clavarioid basidiocarps and lichenised nutritional mode (Petersen 1967, Fischer et al. 2007, Nelsen et al. 2007, Masumoto & Degawa 2020b. Multiclavula is morphologically and ecologically similar to another lichenised genus Bryoclavula but it has a globular or bulbil-like structure which is lacking in Bryoclavula and phylogenetically it is distant from Bryoclavula (Masumoto & Degawa 2020a). ...
... Multiclavula is morphologically and ecologically similar to another lichenised genus Bryoclavula but it has a globular or bulbil-like structure which is lacking in Bryoclavula and phylogenetically it is distant from Bryoclavula (Masumoto & Degawa 2020a). Multiclavula consists of thirteen species from Asia, Europe, North America, Oceania and South America (Corner 1950, Petersen 1967, Petersen 1988, Nelsen et al. 2007, Masumoto & Degawa 2020b. ...
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... It is a lichen-forming basidiomycete, that together with M. vernalis, are the only basidiolichens currently known to occur in southern Ontario. The lichenized thallus is composed of small bulbils of algal cells surrounded by a thick layer of fungal hyphae (Nelsen et al. 2007, Oberwinkler 1984), and the thallus covers the substrate from which fruiting bodies (basidiocarps) are occasionally produced. Because the cream-coloured or yellowish basidiocarps are not perennial, records of M. mucida are scarce. ...
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... Increasing focus was also placed on species recognition in lichenized Basidiomycota, using the fungal ITS barcoding locus (Schoch et al. 2012;Lücking et al. 2014b). As a result, the number of accepted species of lichenized Basidiomycota, including those described herein, has increased dramatically, in particular in the genus Cora (Fig. 1) (Chaves et al. 2004;Fischer et al. 2007;Nelsen et al. 2007;Hodkinson et al. 2014;Sulzbacher et al. 2012;Yánez et al. 2012;Lücking et al. 2013aLücking et al. , b, 2014bSchmull et al. 2014;Vargas et al. 2014;Yanaga et al. 2015;Dal Forno et al. 2016). This corresponds to an increase of the proportion of Basidiomycota among lichen fungi from approximately 0.2 % until the year 2000 to 0.9 % currently and 1.6 % predicted by the end of 2017. ...
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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, Madriñán & Lücking spec. nov., C. applanata Moncada, Soto-Medina & Lücking spec. nov., C. arachnodavidea Moncada, Dal Forno & Lücking spec. nov., C. arborescens Dal Forno, Chaves & Lücking spec. nov., C. arcabucana Moncada, C. Rodríguez & Lücking spec. nov., C. aturucoa Lücking, Moncada & C. Vargas spec. nov., C. auriculeslia Moncada, Yánez-Ayabaca & Lücking spec. nov., C. barbifera Moncada, Patiño & Lücking spec. nov., C. boleslia Lücking, 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 & Lücking spec. nov., C. caraana Lücking, Martins & Lucheta spec. nov., C. casasolana Moncada, R.-E. Pérez & Lücking spec. nov., C. caucensis Moncada, M. Gut. & Lücking spec. nov., C. celestinoa Moncada, Cabrera-Amaya & Lücking spec. nov., C. comaltepeca Moncada, R.-E. Pérez & Herrera-Camp. spec. nov., C. corani Lücking, E. Morales & Dal Forno spec. nov., C. corelleslia Moncada, A. Suárez-Corredor & Lücking spec. nov., C. crispoleslia Moncada, J. Molina & Lücking spec. nov., C. cuzcoensis Holgado, Rivas Plata & Perlmutter spec. nov., C. dalehana Moncada, Madriñán & Lücking spec. nov., C. davibogotana Lücking, Moncada & Coca spec. nov., C. davicrinita Moncada, Madriñán & Lücking spec. nov., C. davidia Moncada, L. Vargas & Lücking spec. nov., C. dewisanti Moncada, A. Suárez-Corredor & Lücking spec. nov., C. dulcis Moncada, R.-E. Pérez & Lücking spec. nov., C. elephas Lücking, Moncada & L. Vargas spec. nov., C. fuscodavidiana Lücking, Moncada & L. Vargas spec. nov., C. garagoa Simijaca, Moncada & Lücking spec. nov., C. gigantea Lücking, Moncada & Coca spec. nov., C. gomeziana Dal Forno, Chaves & Lücking spec. nov., C. guajalitensis Lücking, Robayo & Dal Forno spec. nov., C. hafecesweorthensis Moncada, Lücking & R. Peláez spec. nov., C. haledana Dal Forno, Chaves & Lücking spec. nov., C. hawksworthiana Dal Forno, P. Nelson & Lücking spec. nov., C. hochesuordensis Lücking, E. Morales & Dal Forno spec. nov., C. hymenocarpa Lücking, Chaves & Lawrey spec. nov., C. imi Lücking, Chaves & Lawrey spec. nov., C. itabaiana Dal Forno, Aptroot & M. Cáceres spec. nov., C. leslactuca Lücking, Moncada & R. Peláez spec. nov., C. maxima Wilk, Dal Forno & Lücking spec. nov., C. minutula Lücking, Moncada & Yánez-Ayabaca spec. nov., C. palaeotropica Weerakoon, Aptroot & Lücking spec. nov., C. palustris Dal Forno, Chaves & Lücking spec. nov., C. parabovei Dal Forno, Kukwa & Lücking spec. nov., C. paraciferrii Lücking, Moncada & J.E. Hern. spec. nov., C. paraminor Dal Forno, Chaves & Lücking spec. nov., C. pastorum Moncada, Patiño & Lücking spec. nov., C. pichinchensis Paredes, Jonitz & Dal Forno spec. nov., C. pikynasa J.-M. Torres, Moncada & Lücking spec. nov., C. pseudobovei Wilk, Dal Forno & Lücking spec. nov., C. pseudocorani Lücking, E. Morales & Dal Forno spec. nov., C. putumayensis L.J. Arias, Moncada & Lücking spec. nov., C. quillacinga Moncada, F. Ortega & Lücking spec. nov., C. rothesiorum Moncada, Madriñán & Lücking spec. nov., C. rubrosanguinea Nugra, Moncada & Lücking spec. nov., C. santacruzensis Dal Forno, Bungartz & Yánez-Ayabaca, spec. nov., C. schizophylloides Moncada, C. Rodríguez & Lücking spec. nov., C. smaragdina Lücking, Rivas Plata & Chaves spec. nov., C. soredavidia Dal Forno, Marcelli & Lücking spec. nov., C. subdavicrinita Moncada, J. Molina & Lücking spec. nov., C. suturifera Nugra, Besal & Lücking spec. nov., C. terrestris Dal Forno, Chaves & Lücking spec. nov., C. terricoleslia Wilk, Dal Forno & Lücking spec. nov., C. udebeceana Moncada, R. Peláez & Lücking spec. nov., C. urceolata Moncada, Coca & Lücking spec. nov., C. verjonensis Lücking, Moncada & Dal Forno spec. nov., C. viliewoa Lücking, Chaves & Soto-Medina spec. nov., and C. yukiboa Mercado-Díaz, Moncada & Lücking spec. nov. Furthermore, the taxonomic status of the recently described or recognized species C. arachnoidea, C. aspera, C. ciferrii, and C. reticulifera, is revised.
... Recent investigations indicate that the Dictyonema clade might be a source of a remarkable number of undescribed species (Lücking et al. 2013a(Lücking et al. , 2014b. Studies have revealed a previously unrecognized diversity of basidiolichens in a variety of phylogenetic groups, not only in the Dictyonema clade (Chaves et al. 2004;Yánez et al. 2012;Dal-Forno et al. 2013;Lücking et al. 2013aLücking et al. , b, 2014b but also in the chlorolichen genera Lichenomphalia (Redhead et al. 2002), Lepidostromatales (Fisher et al. 2007;Ertz et al. 2008;Hodkinson et al. 2012Hodkinson et al. , 2014Sulzbacher et al. 2013;Yanaga et al. 2015) and Multiclavula (Nelsen et al. 2007). Our hypothesis is that Acantholichen also may represent a potentially important source of new species that warrants further investigation. ...
... Recent investigations indicate that the Dictyonema clade might be a source of a remarkable number of undescribed species (Lücking et al. 2013a(Lücking et al. , 2014b. Studies have revealed a previously unrecognized diversity of basidiolichens in a variety of phylogenetic groups, not only in the Dictyonema clade (Chaves et al. 2004;Yánez et al. 2012;Dal-Forno et al. 2013;Lücking et al. 2013aLücking et al. , b, 2014b but also in the chlorolichen genera Lichenomphalia (Redhead et al. 2002), Lepidostromatales (Fisher et al. 2007;Ertz et al. 2008;Hodkinson et al. 2012Hodkinson et al. , 2014Sulzbacher et al. 2013;Yanaga et al. 2015) and Multiclavula (Nelsen et al. 2007). Our hypothesis is that Acantholichen also may represent a potentially important source of new species that warrants further investigation. ...
... Most basidiolichens are concentrated in the family Hygrophoraceae (Agaricales, Agaricomycetidae), which includes a great variety of basidiome types, including agaricoid, cyphelloid, stereoid and corticioid (Lawrey et al. 2009;Dal-Forno et al. 2013;Lodge et al. 2014). Basidiolichens with clavarioid basidiomes are known from two orders, Cantharellales and Lepidostromatales (Nelsen et al. 2007;Ertz et al. 2008;Hodkinson et al. 2014). ...
... Although the number of basidiolichens was believed to be relatively small, recent efforts have led to an increase in the diversity of this group of organisms through the discovery of new species in several countries, especially in the Neotropics, such as Costa Rica, Ecuador and Brazil (Chaves et al. 2004;Nelsen et al. 2007;Sulzbacher et al. 2012;Yánez et al. 2012;Dal-Forno et al. 2013;Lücking et al. 2013Lücking et al. , 2014aHodkinson et al. 2014;Schmull et al. 2014). ...
... One possible character to partially separate both genera is the number of sterigmata. Multiclavula generally has 4-8 sterigmata, as frequently observed in other taxa in Cantharellales (e.g., Petersen 1967;Moncalvo et al. 2006;Nelsen et al. 2007), while most other basidiomycetes, including Lepidostromatales, have (1-)2-4 sterigmata (Oberwinkler 1984;Fischer et al. 2007;Ertz et al. 2008;Hodkinson et al. 2012;Sulzbacher et al. 2012). According to Petersen (1967) (Petersen 1967) and some of these might represent Lepidostromatales, although M. vernalis has been confirmed in Multiclavula using sequence data. ...
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Sulzbacheromyces is a recently erected genus in Lepidostromatales, differing from Lepidostroma in the crustose thallus. After the initial discovery of S. caatingae, the only species to be found in Brazil so far, a large quantity of additional data and ITS barcoding sequences for this taxon from a much broader geographical range and different habitats was collected. Phylogenetic analysis under a maximum likelihood framework demonstrated that all specimens are genetically uniform, showing no variation in their ITS, suggesting that S. caatingae has a wide ecological amplitude beyond the Brazilian Caatinga and Atlantic Forest biomes. Detailed descriptions and illustrations of the species are presented, including a map showing the distribution of S. caatingae in the Brazilian semi-arid region and the north-eastern Atlantic rainforest.