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Habit images of the thallus and ascomata from select Trypetheliaceae taxa. A, Arthopyrenia cinchonae (Costa Rica, Lücking 45 ); B, Mycomicrothelia hemisphaerica (Costa Rica, Lücking s.n. ); C, Pseudopyrenula subnudata (Fiji, Lumbsch 19845g ); D, Polymeridium cata- pastum (Bahamas, Britton 6650 ); E, Trypethelium tropicum (Colombia, Lücking 32544 ); F, Campylothelium puiggarii (Brazil, Osorio SM25 ); G, Marcelaria purpurina (Colombia, Moncada 3467 ); H, Trypethelium eluteriae (Colombia, Moncada 3399 ); I, Laurera sanguinaria (Brazil, Brako 7136 ); J, Aptrootia elatior (New Zealand, Walker s.n. ); K, Architrypethelium seminudum (Costa Rica, Lücking 15212b ); L, Astrothelium galbineum (Florida, Harris 41750 ); M, Trypethelium nitidiusculum (Costa Rica, Lücking 34530 ); N, Cryptothelium rhodotitthon (Brazil, Dumont 596 ); O, Laurera megasperma (Costa Rica, Lücking s.n. ). — Scale bars = 1 mm. All images by R. Lücking. 

Habit images of the thallus and ascomata from select Trypetheliaceae taxa. A, Arthopyrenia cinchonae (Costa Rica, Lücking 45 ); B, Mycomicrothelia hemisphaerica (Costa Rica, Lücking s.n. ); C, Pseudopyrenula subnudata (Fiji, Lumbsch 19845g ); D, Polymeridium cata- pastum (Bahamas, Britton 6650 ); E, Trypethelium tropicum (Colombia, Lücking 32544 ); F, Campylothelium puiggarii (Brazil, Osorio SM25 ); G, Marcelaria purpurina (Colombia, Moncada 3467 ); H, Trypethelium eluteriae (Colombia, Moncada 3399 ); I, Laurera sanguinaria (Brazil, Brako 7136 ); J, Aptrootia elatior (New Zealand, Walker s.n. ); K, Architrypethelium seminudum (Costa Rica, Lücking 15212b ); L, Astrothelium galbineum (Florida, Harris 41750 ); M, Trypethelium nitidiusculum (Costa Rica, Lücking 34530 ); N, Cryptothelium rhodotitthon (Brazil, Dumont 596 ); O, Laurera megasperma (Costa Rica, Lücking s.n. ). — Scale bars = 1 mm. All images by R. Lücking. 

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While the phylogenetic position of Trypetheliaceae has been the subject of recent molecular studies, the relationships within this family have been little studied. Here we construct a detailed genus-level phylogeny of the family. We confirm previous morphology-based findings suggesting that a substantial proportion of genera are not monophyletic, a...

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... in the Trypethelium eluteriae Sprengel group, i.e., Trypethelium s.str. (including T. eluteriae, T. platystomum Mont. and T. inamoenum Müll.Arg.) formed a strongly supported clade. Species in this group frequently produce ascomata in pseudostroma, which usually contain, and are often also covered with anthraquinone pigments (Fig. 1H). In addition, these species produce thin-walled ascospores with oval to rectangular lumina (Fig. 2J). Sister to the T. eluteriae group is a clade representing the recently segregated Marcelaria, composed of M. purpurina (Nyl.) Aptroot & al. (Fig. 1G) and M. cumingii (Mont.) Aptroot & al., which have traditionally been placed in ...
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... in pseudostroma, which usually contain, and are often also covered with anthraquinone pigments (Fig. 1H). In addition, these species produce thin-walled ascospores with oval to rectangular lumina (Fig. 2J). Sister to the T. eluteriae group is a clade representing the recently segregated Marcelaria, composed of M. purpurina (Nyl.) Aptroot & al. (Fig. 1G) and M. cumingii (Mont.) Aptroot & al., which have traditionally been placed in Laurera. These species produce ascomata with a broad ostiole, as well as large muriform ...
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... species formed an unsupported, monophyletic group. These taxa produce ascomata with a lateral ostiole (Fig. 1F) and hyaline, muriform ascospores. Trypethelium virens Tuck. and an undescribed taxon, both of which produce hyaline, transversely septate ascospores, formed a strongly supported group together with Campylothelium species, even if clearly differing morphologically. Relationships within this clade remain obscure, due to low internal ...
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... a large number of taxa grouped in the "Astrothelium" clade, which was composed of species currently placed in Astrothelium, Bathelium, Cryptothelium, Laurera and Trypethelium (Fig. 1L-O). Species from most of these genera do not form monophyletic groups within this clade but instead appear mixed. This clade is composed of taxa with variable ascomatal morphology and hyaline ascospores that are muriform or transversely septate (Figs. ...
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... species currently placed in Astrothelium, Bathelium, Cryptothelium, Laurera and Trypethelium (Fig. 1L-O). Species from most of these genera do not form monophyletic groups within this clade but instead appear mixed. This clade is composed of taxa with variable ascomatal morphology and hyaline ascospores that are muriform or transversely septate (Figs. ...
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... "Astrothelium" clade is pantropical and composed of corticolous taxa, producing corticate thalli with ascomata embedded in pseudostromata or thalline warts/galls (Fig. 1L-O). A large amount of variability exists in the ascomatal arrangement (solitary vs. aggregated vs. fused), ostiole orientation (apical vs. eccentric), ascospore septation (transversely septate vs. muriform) and size. All taxa produce hyaline ascospores; transversely septate ascospores are of the astrothelioid-type, which are characterized ...
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... the T. eluteriae group is closely related to Marcelaria (Laurera purpurina and L. cumingii groups). As T. eluteriae is the type of the genus, this clade represents Trypethelium s.str. This group is comprised of several corticolous species that produce ascomata aggregated in large pseudostroma that usually contain brightly colored anthraquinones (Fig. 1H). In addition, this group produces transversely septate ascospores with a reduced endospore forming rounded to oval lumina (Fig. 2J), which are different from the diamond-shaped lumina produced by most other species in the family (Fig. 2C). However, ascospores in T. eluteriae pass through an astrothelioid stage early in their ontogeny ...
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... three species of Aptrootia form a strongly supported monophyletic group sister to the Architrypethelium + "Astrothelium" clade. Aptrootia species produce thin, cartilaginous and bullate or verrucose thalli (Fig. 1J), and form one to two, extremely large, dark brown, muriform ascospores per ascus (Fig. 2M-O), which pass through an initial astrothelioid stage during their ontogeny (Sweetwood & al., 2012). These taxa typically grow at high altitudes on soil, although A. elatior (Stirt.) Aptroot has also been found on bark; consequently, Aptrootia ...
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... is a small, pantropical, corticolous genus producing solitary ascomata with lateral ostioles and muriform ascospores (Fig. 1F). This genus differs from Laurera primarily in its production of lateral ostioles. Harris (1995) and argued that many of the genera found in the "Astrothelium" clade would contain species of the genus Campylothelium. In the present study, the Campylothelium species included were placed outside the "Astrothelium" clade and in a weakly ...
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... "Astrothelium" clade would contain species of the genus Campylothelium. In the present study, the Campylothelium species included were placed outside the "Astrothelium" clade and in a weakly supported monophyletic group. However, only two previously described species were included in our study, and one of these, Campylothelium puiggarii Müll.Arg. (Fig. 1F), which is the type of Campylothelium, has previously been regarded as an unusual species in the genus, with Harris (1995) suggesting it would be placed outside the large "Astrothelium" clade. Sampling of additional Campylothelium species may reveal their placement in the "Astrothelium" clade. Trypethelium virens and an undescribed ...
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... Marcelaria purpurina ( Fig. 1G) and M. cumingii were sister to the T. eluteriae group. This genus was recently described to accommodate Laurera purpurina (Nyl.) Zahlbr., L. cumingii (Mont.) Zahlbr., and L. benguelensis (Müll. Arg.) Zahlbr. (Aptroot & al., 2013b). Marcelaria purpurina, the type species of the genus, produces large ascomata that are bright red. The ...
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... further genus recovered as monophyletic, including the type, P. diluta, was the pantropical and corticolous Pseudopyrenula (Fig. 1C). Pseudopyrenula forms an ecorticate thallus with solitary ascomata with hyaline, three-septate ascospores. Aptroot (1991a) suggested Pseudopyrenula occupied a basal position in Trypetheliaceae, and somewhat similarly, Harris (1998) suggested Pseudopyrenula represented an extreme in the family; however, it was unclear to Harris which ...
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... type, photobiont association and lack of mesospore thickening. This section was later raised to the genus level (Harris in Tucker & Harris, 1980) and has been thoroughly treated in Harris (1991) and Aptroot & Cáceres (2014); an additional, moleculer study verified its inclusion in Trypetheliaceae (Nelsen & al., 2011b). Polymeridium species (Fig. 1D) are tropical, drought tolerant, high-light requiring, and relatively fast-growing taxa that occur mostly in dry forests such as the Brazilian Caatinga (Harris, 1991;Cáceres, 2007). Most species are bark-inhabiting, although a bambusicolous taxon, Polymeridium bambusicola Aptroot & Ferraro, is known (Aptroot & Ferraro, 2000). In the ...
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... cannot be confirmed here except that both diverge relatively early within the tree, although support is weak in critical parts of the tree bearing on this hypothesis. Two species of Polymeridium, P. albocinereum and P. catapastum, formed a supported relationship with Trypethelium tropicum. Harris (1984) had suggested that Trypethelium tropicum (Fig. 1E) produced a thallus similar to other Trypethelium species, but its ascomata were similar to those of Pseudopyrenula. The ascomata of T. tropicum are rather unusual among Trypetheliaceae taxa, producing a barrel-shaped fruiting body with a flat top. Our data confirm that T. tropicum is neither closely related to Trypethelium, nor to ...
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... both suggested a close relationship between Mycomicrothelia and Arthopyrenia s.str., with Hawksworth (1985) noting similarities between the two genera, such as the production of partially exposed ascomata and pycnidia containing bacillariform conidia (which are present in many Mycomicrothelia species). Hawksworth (1985) noted that Mycomicrothelia (Fig. 1B) species differ from Arthopyrenia (Fig. 1A) in the verruculose ornamentation of their ascospores, and also in that ascospores of Mycomicrothelia species turn brown while inside the asci. Aptroot (1995) suggested a probable placement in Arthopyreniaceae. Therefore, after finding Mycomicrothelia species belong to Trypetheliaceae (Nelsen & ...
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... Mycomicrothelia and Arthopyrenia s.str., with Hawksworth (1985) noting similarities between the two genera, such as the production of partially exposed ascomata and pycnidia containing bacillariform conidia (which are present in many Mycomicrothelia species). Hawksworth (1985) noted that Mycomicrothelia (Fig. 1B) species differ from Arthopyrenia (Fig. 1A) in the verruculose ornamentation of their ascospores, and also in that ascospores of Mycomicrothelia species turn brown while inside the asci. Aptroot (1995) suggested a probable placement in Arthopyreniaceae. Therefore, after finding Mycomicrothelia species belong to Trypetheliaceae (Nelsen & al., 2009), the inclusion of Arthopyrenia ...

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... Ambiguously aligned positions were corrected manually. Additional mtSSU sequences belonging to different genera of Trypetheliaceae were downloaded from GenBank (Table 1), their selection based on the present phylogenetic classification of the family (Del Prado et al. 2006;Nelsen et al. 2009Nelsen et al. , 2014Luangsuphabool et al. 2016Luangsuphabool et al. , 2018Lücking et al. 2016). For the final analysis, we used a subset of selected species from the genera Aptrootia, Architrypethelium, Astrothelium, Bathelium, Pseudopyrenula and Trypethelium. ...
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... The latter genus fell into two clades (Lücking et al. 2016) with one being a sister group to Architrypethelium. Phenotypically Architrypethelium differs from Astrothelium in having predominantly large ascospore without diamond-shaped lumina when mature (Aptroot 1991, Aptroot et al. 2008, Nelsen et al. 2014, Aptroot and Lücking 2016, Lücking et al. 2016b). Another genus with muriform ascospores is Aptrootia, which also shares an astrothelioid stage in the young ascospores (Lücking et al. 2016) and the genus formed a sistergroup to a clade including Architrypethelium and Astrothelium p.pt. further calling the generic delimitation in the family in question. ...
... Morphologically, Aptrootia differs from Astrothelium in having dark brown ascospores with a hard outer shell (Lücking et al. 2016). While most genera in Trypetheliaceae, such as Astrothelium s.str., Bathelium, Polymeridium and Viridothelium include species with various ascospore types (Hyde et al. 2013, Nelsen et al. 2014, Aptroot and Lücking 2016, Lücking et al. 2016b, the species of Architrypethelium shared a similar ascospore morphology (Nelsen et al. 2014, Lücking et al. 2016b. ...
... Morphologically, Aptrootia differs from Astrothelium in having dark brown ascospores with a hard outer shell (Lücking et al. 2016). While most genera in Trypetheliaceae, such as Astrothelium s.str., Bathelium, Polymeridium and Viridothelium include species with various ascospore types (Hyde et al. 2013, Nelsen et al. 2014, Aptroot and Lücking 2016, Lücking et al. 2016b, the species of Architrypethelium shared a similar ascospore morphology (Nelsen et al. 2014, Lücking et al. 2016b. ...
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Architrypetheliummurisporum Luangsuphabool, Lumbsch & Sangvichien is described for a crustose lichen occurring in dry evergreen forest in Thailand. It is characterised by a green to yellow-green corticated thallus, perithecia fused in black pseudostromata with white rim surrounding the ostiole and small, hyaline and muriform ascospores. Currently, all species in the genus Architrypethelium have transversely septate ascospores, hence the discovery of this new species indicates that ascospore septation is variable within the genus, similar to numerous other groups of lichen-forming ascomycetes. Phylogenetic analyses of two loci (mtSSU and nuLSU) supported the position of the new species within Architrypethelium . This is the first report of the genus in Southeast Asia.