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        A revisionary synopsis of the Trypetheliaceae (Ascomycota: Trypetheliales)
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        A revisionary synopsis of the Trypetheliaceae (Ascomycota: Trypetheliales)
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        A revisionary synopsis of the Trypetheliaceae (Ascomycota: Trypetheliales)
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Abstract

A revisionary synopsis is presented for the family Trypetheliaceae, 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), and Viridothelium (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 genera Astrothelium, Campylothelium, Cryptothelium, and Trypethelium, 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 genera Aptrootia (large, brown, muriform ascospores), Architrypethelium (large, mostly 3-septate ascospores), and Pseudopyrenula (ecorticate, white thalli and astrothelioid ascospores) are maintained, Trypethelium is redefined to include species with raised, pseudostromatic ascomata and multiseptate ascospores with thin septa. The sister group of Trypethelium is the genus Marcelaria, with brightly coloured pseudostromata and muriform ascospores. Bathelium is 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 to Arthopyrenia, Mycomicrothelia and Polymeridium, viz. Bogoriella, Constrictolumina, Dictyomeridium, and Novomicrothelia. In addition, separate genera are accepted for the Trypethelium tropicum (Nigrovothelium) and T. 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., and V. megaspermum (Makhija & Patw.) Aptroot & Lücking comb. nov. In addition, six replacement names are proposed: Astrothelium campylocartilagineum Aptroot & Lücking nom. nov., A. grossoides Aptroot & Lücking nom. nov., A. octosporoides Aptroot & Lücking nom. nov., A. scoriothelium Aptroot & Lücking nom. nov., A. pyrenastrosulphureum Aptroot & Lücking nom. nov., and Bathelium pruinolucens Aptroot & 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 in Trypetheliaceae but previously excluded from the family, and their current names, is also provided.

Footnotes

The original version of this article was published with a half-couplet missing from page 964 of the article. A notice detailing this has been published online and the error rectified in the online PDF and HTML copies.

Introduction

The family Trypetheliaceae is an almost strictly tropical lineage of nearly exclusively corticolous (very rarely saxicolous), lichenized fungi (Fig. 1). The first members of this family were encountered towards the end of the 18th century on pieces of medicinal bark (mainly Cinchona for quinine) that had been collected in South American forests (Zenker 1827). The conspicuous, often brightly coloured ascomata with complex structures intrigued lichenologists of the time and both Acharius (1817 a , b ) and Fée (1831) wrote monographs of what was then known as the genus Trypethelium, treating 10 and 19 species, respectively. More species became known especially from Australia, Brazil and Borneo, which were described in separate papers, mostly by Eschweiler (1824, 1833), Krempelhuber (1875), Montagne (1851, 1856) and Müller (1884, 1885). During this time, a further synopsis of the family was made by Trevisan (1861).

Fig. 1 Mosaic of six Astrothelium species (and one Phaeographis) on a tropical rainforest tree at Los Amigos Biological Station in Amazonian Peru. Left, with orange pseudostromata, A. kunzei; centre, with white pseudostromata, A. rufescens; upper right, with green thallus, A. cf. cecidiogenum; right, mottled-green-yellow thallus, A. subscoria; lower right, small thallus, A. nitidiusculum; lower middle and left, A. aeneum.

After this initial burst of activity, which concluded with Malme’s treatment of species gathered during the first Regnellian Expedition in Brazil (Malme 1924), relatively few species were added for a long time, and for many species only the type was known. In the meantime, the genus Trypethelium had been split into smaller entities and various classification systems had been proposed and applied, particularly by Müller (1884, 1885), Nylander (1863), Vainio (1890) and Zahlbruckner (1922, 1924, 1928). These studies doubled the number of names accepted in the Trypetheliaceae solely through reclassifications. Halfway through the 20th century, this led to the situation where almost as many names existed in the family as there had been specimens collected and illustrated by the only monograph published around that time, by Letrouit-Galinou (1957, 1958), on the genus Laurera. In part, the genera of Trypetheliaceae were also mixed with those of Pyrenulaceae since the overall ascoma morphology was considered more important than anatomical features such as hamathecium structure and ascospore type (e.g. Dodge 1953). Both these character complexes clearly separate the two families, which are not even closely related; molecular phylogenetic studies placed Pyrenulaceae in Eurotiomycetes and Trypetheliaceae in Dothideomycetes (Del Prado et al. 2006; Nelsen et al. 2009; Schoch et al. 2009 a , b ).

Aside from two papers published by Johnson (1940, 1959) on North American Trypetheliaceae and an anatomical study of the endoperidermal thallus of Trypethelium eluteriae (Lambright & Tucker 1980), serious taxonomic and systematic treatments of the Trypetheliaceae were only taken up again by Harris with his studies on Amazonian species (Harris 1986) and a revision of Polymeridium (Harris 1993). Previously, Eriksson (1981) had briefly treated the family in his conspectus of bitunicate Ascomycetes and elaborated a detailed evolutionary scheme for the supposedly closely related Trypetheliaceae and Pyrenulaceae. The work by Harris (1986, 1993) redefined the family in a restricted sense and adopted a generic division based on thallus type, ascoma disposition, and ascospore type that has been in use since then. However, Harris (1995) and other workers (Aptroot et al. 2008) also recognized that this system was at least in part artificial, resulting in cases such as in the Astrothelium variolosum-Trypethelium nitidiusculum complex where a single specimen could be partly assigned to one genus and partly to another (Harris 1995). Further systematic treatments were provided by Aptroot (1991), introducing the new genus Architrypethelium, and Harris (1998), with a revision of the genus Pseudopyrenula. Harris’s work in particular triggered further inventories and taxonomic studies of the family, particularly in India but also in the Neotropics and Australia, describing many new taxa (Upreti & Singh 1987; Makhija & Patwardhan 1988, 1992, 1993; McCarthy & Kantvilas 1993; McCarthy 1995; Aptroot & Ferraro 2000; Aptroot et al. 2008; Aptroot 2009 a , b ). At the time when Harris (1986) published his Amazonian studies, Hawksworth (1986) provided a revision of the genus Mycomicrothelia, a genus at the time not thought to be related to Trypetheliaceae but recently found to include many species falling in lineages at the base of the family (Nelsen et al. 2009, 2011, 2014; Lücking et al. 2016 a ) and now redispositioned in the genera Bogoriella and Novomicrothelia (Lücking et al. 2016 a ).

Notably, the Trypetheliaceae were the subject of several detailed studies about ascospore anatomy and ontogeny (Sweetwood et al. 2012) and pigment chemistry (Mathey & Hoder 1978; Mathey 1979; Mathey et al. 1980), as well as potential pharmaceutical properties of their chemical compounds (Manojlovic et al. 2010). Ecological and ecogeographical papers with details on Trypetheliaceae were published by Komposch & Hafellner (2000) on canopy and savanna species in Venezuela, Aptroot et al. (2008) on pyrenocarpous lichens in Costa Rica, Rivas Plata et al. (2008) on the correlation between family-level diversity and environmental parameters related to the conservation status of tropical forest ecosystems, and Aptroot (2009 b ) on diversity and endemism of Trypetheliaceae and Pyrenulaceae in Malaysia.

The systematics of the family regained interest with the advent of molecular phylogenetic methods. An initial study by Del Prado et al. (2006) demonstrated the placement of Trypetheliaceae within Dothideomycetes, and phylogeny of the family was refined through subsequent work by Nelsen et al. (2009, 2011, 2014) and Lücking et al. (2016 a ), showing that lichenized species of Arthopyrenia and Mycomicrothelia also belong here, and laying the groundwork for a much revised genus and species concept (Aptroot et al. 2013 a ; Hyde et al. 2013; Aptroot & Cáceres 2014; Nelsen et al. 2014; Luangsuphabool et al. 2016; Lücking et al. 2016 a ). This also stimulated further inventory work in several parts of the tropics, with the discovery of many new taxa (Aptroot & Cáceres 2013, 2016; Aptroot et al. 2013 b , 2016 a , b ; Lima et al. 2013; Córdova-Chávez et al. 2014; Weerakoon & Aptroot 2014; Flakus et al. 2016; Luangsuphabool et al. 2016; Lücking et al. 2016 b ). The order Trypetheliales was eventually established for the family (Aptroot et al. 2008; Hyde et al. 2013), and recently a second family, Polycoccaceae, including lichenicolous fungi, was included in this order (Ertz et al. 2015).

In the present work, we provide a concise synopsis of the family Trypetheliaceae to account for the numerous changes that have taken place based on recent phylogenetic studies and a new assessment of the delineation of genera and species within the family. Together with the other papers included within this special issue, this should give a complete overview of the current knowledge of the family. We realize that this synopsis cannot be a definitive monographic treatment; the gaps in knowledge about this megadiverse group of tropical lichens are too large and still many taxa are known only from few or single collections. While available sequence data and revision of phenotypic characters suggest that the concept presented here reflects genus and species boundaries more accurately than previous treatments, we expect that this synopsis will also trigger many further studies. These, we anticipate, would be based on a much more thorough sampling of individual geographical regions, which will unearth novel taxa and also require further revision of Trypetheliaceae species. Furthermore, additional types of old names to be assigned to this family will undoubtely surface.

Material and Methods

Identification and descriptive work was carried out in Soest, using an OLYMPUS SZX7 stereomicroscope and an OLYMPUS BX50 compound microscope with interference contrast, connected to a NIKON Coolpix digital camera, as well as at the Field Museum in Chicago, using a LEICA MS5 dissecting microscope and a ZEISS Axioskop 2 compound microscope, in part connected to JENOPTIK ProgRes C3 and C5 digital microscope cameras. Further digital images were taken with CANON Powershot SX20IS, NIKON F301 and Sony Alpha 33 DSLR digital cameras.

Sections of thalli and ascomata were cut by hand with a razor blade and examined with squash preparations on material mounted in water, KOH and Lugol’s solution. All measurements are given for water mounts. Illustrated thin sections were made by Dr Felix Schumm using a Wild M3 stereomicroscope, an Olympus BX51 compound microscope with interference contrast, a Canon EOS 40D camera with MP-E 65 mm and a Mic HM 560 cryotome. Specimens were measured and illustrated in tap water, unless marked with IKI (mounted in iodine) or LCB (mounted in lactophenol cotton blue).

Chemistry was analyzed by means of a UV lamp and K-tests using 10% KOH solution, both on thallus and ascoma portions including medulla and on sections. Many specimens, including most types (where not previously analyzed), were also investigated using thin-layer chromatography (TLC) in solvents A and/or C (Orange et al. 2010).

Well over 2000 specimens were examined for this study, including most types. Where possible, types were studied through loans or visits to the corresponding major herbaria; in some cases we relied on digitized type specimens available through the Global Plants Initiative (GPI) at JSTOR (https://plants.jstor.org). To complete our own 352 images of type specimens accumulated through studies over the past decades, we added 54 partial images of selected type specimens posted on JSTOR to the plates published here to highlight specific morphological features required for taxon identification, reproduced with permission from the curators at BM (Figs 31E, 34H, 41L, 44F, 49F, 51I, 52K), G (Figs 13F, 15B, 16J & K, 18C & K, 19G, 23C & E, 26G, 28J, 29B & F, 32H, 33B, 35B & I, 37A, G & H, 39G, 44K & L, 52F, 53I, 54F & G, 57G), H (Figs 13D & I, 15H, 18G, 20E, 21A, 27F, 35H, 38I, 43A, B & J, 44B, 48A, 49G), and S (Figs 31D, 39I, 42H, 54A). Quite a number of additional type species not shown here are available through GPI at JSTOR and can be consulted there in case of doubts. Except for types, specimens studied are not cited in full; instead, we summarize the known distribution and give selected new country records. Distributions are given in geographical order following the Flora Neotropica style, from NW to SE, starting with North America and ending with Australia and Oceania.

The numerous collections made by Charles Wright in the 19th century in Cuba, which have been studied (and distributed in exsiccatae) by Tuckerman (1864), Nylander (1876), and Müller (1885, 1894) was an important reference. The citation of these exsiccatae or the corresponding collections has been somewhat ambiguous in the literature, usually giving Wright as editor or with the exsiccate number as his collection number. However, Wright did not perform identification work on his lichen collections; instead they were organized and edited by Tuckerman (as Caroli Wright Lichenes Insulae Cubae or Lich. Ins. Cub.) and Müller (as Graphideae Cubenses (a cl. C. Wright lectae et a cl. W. Nylander determinatae) or Graph. Cub., as Lichenes Cubenses (a cl. Wright lecti et a cl. W. Nylandero pr. p. determinati) or Lich. Cub., and as Verrucariae Cubenses or Verr. Cub.) (http://indexs.botanischestaatssammlung.de)) using a numbering system separate from Wright’s collection numbers. Wright’s original collection numbers are usually not available for his Cuban lichen collections (if he numbered them at all) and they will be cited as Wright s. n.

For the general section, a selection of morphological and anatomical images was provided by Felix Schumm and for the taxonomic section we also included images of five new species of Astrothelium described elsewhere in this issue (Luangsuphabool et al. 2016), provided by Ek Sangvichien.

Morphology, Anatomy and Chemistry

Morphological, anatomical and chemical characters in Trypetheliaceae are variable and, as in other species-rich families of tropical lichens such as Graphidaceae and Pyrenulaceae, provide a broad array of features for taxonomic and systematic purposes, which in the past have not been fully explored due to the lack of a phylogenetic framework to assess the validity of individual characters, in particular the external morphology of thallus and ascomata. This framework is now available (Nelsen et al. 2014; Lücking et al. 2016 a ); it shows that morphological variation changes with evolutionary history within the family. The typical and often unique characters of Trypetheliaceae, particularly the astrothelioid ascospores and the thin, net-like hamathecium, often with coarse inspersion, are mostly developed in derived lineages, whereas the more basal lineages are not particularly different from similar, lichenized forms in other, often unrelated orders, such as Monoblastiales, Strigulales and even certain Pyrenulales.

Thallus morphology

Various characters can be assessed with regard to thallus morphology, including whether the thallus is (partially) endoperidermal, the presence of a cortex, the development of surface structures, colour and the associated pigment chemistry (see also below). Basal lineages in the family, such as former species of Arthopyrenia and Mycomicrothelia now dispositioned in the genera Bogoriella, Constrictolumina, and Novomicrothelia, as well as the genera Dictyomeridium, Distothelia, Polymeridium, and Pseudopyrenula, almost invariably have an ecorticate, whitish thallus that is often endoperidermal (Fig. 2A–F), with scattered photobiont cells; further structures except for a black prothallus line (Fig. 2F) are usually absent and the ascomata are usually exposed and black (Fig. 2A–F). Except for Pseudopyrenula, this correlates with the presence of non-astrothelioid ascospores with thin or uniformly thickened septa and rectangular to rounded, but not diamond-shaped, lumina (see below).

Fig. 2 Thallus morphology and anatomy in Trypetheliaceae. A–C, endoperidermal thallus with outline of periderm cells visible; A, Bogoriella modesta; B, Constrictolumina majuscula; C, Bogoriella megaspora. D–F, ecorticate, whitish thallus with black prothallus line; D, B. miculiformis; E, Polymeridium catapastum; F, P. subcinereum. G–I, corticate, (olive-)green thallus; G, Astrothelium megaspermum; H, A. intermedium; I, Nigrovothelium tropicum. J–L, anatomy of epiperidermal thallus showing gelatinized cortex, photobiont layer, and periderm; J & K, Astrothelium tuberculosum; L, Architrypethelium grande. Scales: A–I=1 mm; J–L=20 µm. (Images B, C, J & K by F. Schumm).

In more derived lineages, the thallus is almost invariably corticate and mostly has an olive-green colour when fresh (Fig. 2G–I). The cortex is epiperidermal and often massive, consisting of a loose network of anastomosing hyphae (similar to the hamathecium, but more irregular; see below) embedded in a gelatinous matrix and becoming more dense and orange-yellow in K; the photobiont layer and medulla (if developed) are either epiperidermal or partly endoperidermal (Fig. 2J–L).

While most species have a smooth to uneven thallus (Fig. 3A–C), a particular feature in Trypetheliaceae rarely found in other families such as Graphidaceae, is the formation of a verrucose-papillose to bullate or folded thallus surface (Fig. 3D–I). In section, these structures either include amorphous white crystal masses or mainly consist of a strongly thickened cortex, with anastomosing hyphae embedded in a gelatinous matrix (Fig. 3J & K). In these cases, in which the thallus might even appear squamulose (Fig. 3I), the photobiont layer is often vertically arranged (Fig. 3J & K), suggesting adaptation to high light intensities, for example in the rainforest canopy where such Trypetheliaceae primarily occur. The available phylogenetic data suggest that such thallus structures are species-specific, although habitat-induced variation has been shown in other groups, such as Graphis pseudocinerea in Graphidaceae (Lücking et al. 2011).

Fig. 3 Thallus surface morphology in Trypetheliaceae. A–C, surface smooth to uneven, following the contours of the bark; A, Astrothelium nitidiusculum; B & C, A. rufescens. D, surface papillose-verrucose, A. papillosum. E, surface bullate, A. tuberculosum. F & G, surface bullate-folded; F, A. megalophthalmum; G, A. versicolor. H & I, surface verrucose-bullate-squamulose; H, Aptrootia elatior; I, Astrothelium puiggarii. J & K, section through bullate ‘squamules’ showing gelatinized cortex and vertically arranged photobiont layer, A. puiggarii. L, true gall formation, A. ceratinum. Scales: A–I & L=1 mm; J & K=20 µm. (Images J & K by F. Schumm).

Independent of the surface structure, many Trypetheliaceae seem to produce thickened bark, which suggests gall formation. This is particularly seen in relation to the ascomata, which often emerge from beneath the upper periderm layers of the bark (see below). However, galls in the strict sense should produce abnormal growth of the periderm, in particular the inner, living layers, viz. the cork cambium (phellogen), which due to its meristematic nature is predestined to produce abnormal growth reactions. In most Trypetheliaceae, the endoperidermal thallus portion is developed only in the upper, dead layers of the periderm (Fig. 3J & K), the cork (phellem), and rather than causing growth of the periderm, the thickening appears to be caused by uplifting of the upper layers, particularly above the ascomata (see below). However, some cases have been observed with true gall formation due to growth of the cork (Fig. 3L). Further studies, particularly on the ontogeny of the thallus and ascomata, are needed to properly understand this phenomenon and its biological implications.

Ascoma morphology

The ascomata in Trypetheliaceae provide some of the richest character sets for taxonomic purposes but at the same time have provided stumbling blocks to properly recognizing and classifying the variation encountered in this family. Several characters can be assessed when analyzing ascoma morphology: 1) arrangement of the ascomata relative to the thallus, 2) orientation of the ostioles, 3) ascoma emergence, 4) degree of thallus cover, 5) configuration of the ostiolar area, and 6) pigmentation (see also below).

Ascomata arrangement can vary from regularly dispersed over the thallus (solitary), to confluent, aggregate, and pseudostromatic (Fig. 4A–L), with the ostioles apical, eccentric (lateral) or fused, in which case the shared portion of the ostiole might be apical or lateral (Fig. 5A–L). Ascoma arrangement can best be assessed when comparing distances between individual ascomata. In dispersed ascomata (Fig. 4A–C), random measurement of paired distances will result in a broad, continuous distribution of distance measures. In contrast, aggregate or pseudostromatic ascomata (Fig. 4E–L) will give a bimodal distribution of randomly measured distances, reflecting small distances between ascomata belonging to the same cluster versus large distances between ascomata belonging to different clusters. In specimens with partially confluent ascomata (Fig. 4D), the distribution will be intermediate between the two extremes. To distinguish between aggregate and pseudostromatic clusters, pseudostromata are defined here as areas that differ from the surrounding thallus in either structure (e.g. being strongly emergent; Fig. 4E), colour (often whitish or dominated by black excipular tissue; Fig. 4F–K), or chemistry (e.g. pigment or lichexanthone limited to the pseudostromata; Fig. 4L), or a combination of these. Thus, aggregate ascomata in which the associated thallus is similar to the surrounding thallus and which lack specific features are not considered pseudostromatic. This applies in particular to fused ascomata with a shared ostiole in many species of Astrothelium, in which the thallus associated with the ascomata is not particularly differentiated.

Fig. 4 Ascoma arrangement in Trypetheliaceae. A, solitary, dispersed, Pseudopyrenula subnudata; B, solitary, dense, Astrothelium floridanum; C, solitary, crowded, Nigrovothelium tropicum; D, solitary to confluent, Astrothelium nitidiusculum; E, pseudostromatic by emergence, A. intermedium; F, pseudostromatic by colour contrast, A. norisianum; G, with fused ostioles, fused ascomata dispersed, A. eustomum; H, with fused ostioles, fused ascomata pseudostromatic, A. interjectum; I, pseudostromatic with exposed ascomata and linear-reticulate pseudostromata, A. neogalbineum; J, pseudostromatic with exposed, prominent to sessile ascomata, Bathelium mastoideum; K, pseudostromatic with erumpent to prominent pseudostromata with white cover, Astrothelium sphaerioides; L, pseudostromatic with prominent to sessile pseudostromata covered by pigment, Trypethelium eluteriae. Scales=1 mm.

Fig. 5 Ascoma disposition and orientation of ostioles in Trypetheliaceae. A–C, ostioles apical, separate, A, Polymeridium subcinereum; B, Astrothelium nitidiusculum; C, Trypethelium subeluteriae. D, ostioles apical, with ostiolar areas forming lobate pattern, T. astroideum. E, ostioles apical, ostiolar area separated from covering thallus by a split, Marcelaria purpurina. F–I, ostioles lateral, eccentric, separate, pointing in various directions, F, Astrothelium gigasporum; G, A. scorizum; H, Polymeridium simulans; I, Dictyomeridium proponens. J–L, ostioles lateral, eccentric, centrally fused to form a shared channel leading to various chambers, J, Astrothelium macrocarpum; K, A. marcidum; L, A. intermedium. Scales=1 mm.

For ascoma emergence, we apply here the same criteria and terminology as used in Graphidaceae and the genus Graphis (Lücking 2009), since this feature has often been imprecisely treated in the literature and the term emergent has been used for morphologies ranging from slightly emergent to sessile. Here we define four states: 1) immersed, if more than 3/4 of the ascoma is beneath the thallus level (Fig. 6A); 2) erumpent, if more than 1/4 to 1/2 of the ascoma is above thallus level (Fig. 6B); 3) prominent, if more than 1/2 of the ascoma is above thallus level and the base is expanded outwards to vertical (Fig. 6C); 4) sessile, if more than 3/4 of the ascoma is above thallus level and the base is constricted (Fig. 6D). This terminology also applies to pseudostromata as a whole. A special feature of some Trypetheliaceae is that the ascomata can be deeply immersed, well below the thallus and often deep into the periderm, up to 2 mm and occasionally up to 5 mm below the thallus surface. This is not restricted to a certain taxonomic group but occurs, for example, in Astrothelium (Fig. 6A), Polymeridium, and Pseudopyrenula. Ascomata that are deeply immersed in the periderm are rare in lichens, otherwise mostly known from Pyrenulaceae.

Fig. 6 Ascoma emergence and thallus cover in Trypetheliaceae. A, immersed, Astrothelium calosporum. B, erumpent, A. nitidiusculum. C, prominent, A. megaspermum. D, sessile, A. fallax. E–G, fully exposed, without thallus cover; E, Pseudopyrenula dubia; F, Bathelium porinosporum; G, Astrothelium infuscatulum. H, with basal thalline cover, Polymeridium amyloideum. I, with lateral thalline cover, Astrothelium straminicolor. J, with complete thalline cover, A. megeustomum. K, whitish and exposed ostiolar area, A. fijiense. L, with pigment cover, A. macrocarpum. Scales=1 mm.

With regard to the degree of thallus cover, the following states are distinguished, also based on classifications developed for Graphidaceae and Graphis (Lücking 2009): 1) absent, ascomata are fully exposed from the base (Fig. 6E–G); 2) basal, only the base up to c. 1/4 of the ascoma is covered (Fig. 6H); 3) partial, the ascomata are covered with thallus up to about 3/4 except the area surrounding the ostiole (Fig. 6I); 4) complete, only the ostiole or ostiolar spot remains visible (Fig. 6J). This scheme also applies to pseudostromata or thallus warts with aggregated ascomata (Fig. 6K & L). The configuration of the ostiolar area is also an important taxonomic character, including its visibility as a narrow dot (Fig. 6B) or broad spot (Fig. 6C & D), its colour (mostly black, sometimes white; Fig. 6J), and the presence of a differently coloured, ring-shaped band around the ostiole (usually whitish), which can be sharp and regular (Fig. 6C) or diffuse and often irregular (Fig. 6B & G). In some cases, the entire ascoma is covered with a layer different from the thallus, which is often the case in pseudostromata but can also be found in species with solitary ascomata (Fig. 6K).

Ascoma anatomy

The internal anatomy of the ascomata including the pseudostromata has not been well studied, even though it has the potential to provide useful taxonomic and systematic characters. For instance, Makhija & Patwardhan (1993) made a typology of pseudostroma configurations found in Trypethelium, but more data are needed to fully assess the importance of their categories. Letrouit-Galinou (1957, 1958) also classified the ascomata of the artificial genus Laurera according to wall structures which have subsequently been used, in part, to disperse the species studied over various lineages, including Marcelaria, Bathelium, and Astrothelium. Notably, the species belonging to Marcelaria were placed in different categories by Letrouit-Galinou (1957, 1958), although the main difference is the nature of the pigments present on and in the pseudostromata (Schumm & Aptroot 2012; Aptroot et al. 2013 a ).

The ascomata in Trypetheliaceae develop either superficially above the periderm or emerge from beneath the periderm and while this appears to be species-specific, both epiperidermal and endoperidermal ascomata can be found in different species of the same genus. For example, Architrypethelium grande, Astrothelium megaspermum, A. porosum, A. puiggarii, Marcelaria purpurina, Nigrovothelium tropicum and Trypethelium eluteriae all produce epiperidermal ascomata or pseudostromata, with the periderm remaining below the ascomata and not included in the covering layers (Fig. 7A–F, J–L). In contrast, in Astrothelium tuberculosum, Constrictolumina majuscula and Pseudopyrenula diluta the ascomata are endoperidermal (Fig. 7G–I). The ascoma wall is (1–)2–3-layered with a brown to carbonized excipulum (Fig. 7B–K), often with a cortical layer similar to the thallus cortex, which might be thin (Fig. 7B) to thick (Fig. 7E & F, K & L) and in the case of pseudostromata with an amorphous medullary layer (Fig. 7C); in species such as Astrothelium megaspermum, the upper wall also contains a brownish ‘medullary’ layer which might be homologous to the involucrellum in other pyrenocarpous taxa.

Fig. 7 Ascoma anatomy in Trypetheliaceae. A–C, epiperidermal ascomata with thallus cover; A & B, Astrothelium megaspermum; C, A. porosum. D, epiperidermal ascoma with simple wall, Nigrovothelium tropicum. E–F, epiperidermal ascoma with thick cortical layer, E, Marcelaria purpurina; F, Astrothelium puiggarii. G–I, endoperidermal ascomata, G, A. tuberculosum; H, Pseudopyrenula diluta; I, Constrictolumina majuscula. J, ostiolar area, Trypethelium eluteriae. K–L, gelatinized cortical layer on ascoma, Architrypethelium grande. Scales: A=1 mm; B–K=100 µm; L=50 µm. (Images A-I by F. Schumm).

Ascoma anatomy provides a very rich set of characters but we have not yet fully explored these features here as we await more detailed studies. We expect that ascoma and pseudostroma wall anatomy, explored within a phylogenetic framework, will help to further refine genus and species concepts in Trypetheliaceae.

Hamathecium, asci and ascospores

Most Trypetheliaceae have a rather uniform hamathecium, consisting of thin, straight, much branched and anastomosing paraphyses forming a network embedded in a gelatinous matrix (Fig. 8A). In basal lineages, the paraphyses are usually thicker and less anastomosing and the gelatinous matrix is less obvious. Hymenial inspersion is common throughout the family and usually occurs as large, irregular, colourless oil droplets lining the paraphyses (Fig. 8B) but it can also form a more amorphous, sometimes dirty yellow infusion in basal lineages such as Pseudopyrenula; in some species, the droplets contain anthraquinones and react with K. In most species, inspersion is found along the ostiolar area while the hamathecium is clear. For instance, Harris (1995) suggests muriform-spored species of Bathelium have an inspersed hamathecium, whereas the inspersion in most of these species is developed only around the ostiole. The taxonomic importance of hamathecium inspersion (i.e. of the entire hamathecium) has been neglected, except in a few cases, but phylogenetic studies suggest that it is species-specific (Nelsen et al. 2014; Lücking et al. 2016 a ). Unfortunately most descriptions, especially of historical names, do not mention this character and hence the status of names for which types have not been available is difficult to resolve. On the other hand, inspersion is usually preserved even in old collections as long as hymenium material is present.

Fig. 8 Hamathecium and asci in Trypetheliaceae. A, anatomosing, net-like filaments, Architrypethelium nitens. B, inspersion with oil droplets lining the filaments, Astrothelium megaspermum. C–E, mature ascus with ascospores; C, Constrictolumina majuscula; D, Novomicrothelia oleosa; E, Trypethelium eluteriae. F–K, young and immature asci showing wall layers and apical apparatus; F, Aptrootia terricola; G & H, A. robusta; I, Architrypethelium nitens; J & K, Astrothelium megaspermum. L, ejection of ascospores through ostiole, A. megaspermum. Scales: A–G=20 µm; H–K=100 µm; L=1 mm. (Images C & D by F. Schumm).

Asci in Trypetheliaceae are typically fissitunicate (Eriksson 1981; Aptroot 1991); their internal structure is best observed in species with larger ascospores, as long as the ascospores are young or immature (Fig. 8C–K). Typically the asci have a distinct foot (Fig. 8H & J). The elastic inner wall (endotunica) appears multilayered (Fig. 8J) and the tholus contains a non-staining ring structure. Like ascoma anatomy, ascus structure has not yet been fully studied and explored for taxonomic purposes in this family, but may be helpful to delimit especially the basal lineages.

Besides ascoma morphology, ascospore type is the second most important character complex in Trypetheliaceae (Figs 8L, 9A–L). While in the past ascospore septation was mostly used to separate lineages at the genus level, it is now obvious that the nature of the septa and walls provide the most critical characters at a higher taxonomic level (Fig. 9A–L) whereas septation per se might separate species but varies considerably at genus level. Astrothelioid ascospores are typical of this family , forming secondary wall thickenings that make the lumina diamond-shaped (Fig. 9G). Very similar ascospores occur in some non-lichenized families such as Massariaceae, which Eriksson (1981) used to suggest that they should form part of Trypetheliaceae. In fact these families are not closely related to Trypetheliaceae (Schoch et al. 2009 b ; Hyde et al. 2013). In species of Trypetheliaceae with muriform ascospores, in which the lumina are small and their morphology is difficult to assess, the young ascospores in lineages with an astrothelioid ascospore type undergo a distinctly astrothelioid stage (Fig. 9J & K) (Sweetwood et al. 2012). Almost without exception, a given genus has either astrothelioid or non-astrothelioid ascospores. The latter can be assigned to various types, including the multiseptate ascospores with slightly thickened but not astrothelioid septa in Trypethelium (Fig. 9E) and Viridothelium, or the septate to muriform ascospores in Bathelium (Fig. 9F), Polymeridium (Fig. 9D) and Dictyomeridium. In Architrypethelium the ascospores are principally astrothelioid when immature with a reduced endospore when mature, becoming very large with few septa and often including characteristic, needle-shaped crystals (Fig. 9H). Basal lineages formerly placed in Arthopyrenia and Mycomicrothelia now positioned in the genera Bogoriella, Constrictolumina, and Novomicrothelia have almost invariably 1-septate ascospores with thin or thickened walls and septa but are never astrothelioid (Fig. 9A–D); in Constrictolumina these may form secondary invaginations (partial septa), whereas in Bogoriella and Novomicrothelia, ornamented walls may occur. Distothelia has particular ascospores with strong distal thickenings and the lumina near the central septum (Fig. 9C).

Fig. 9 Ascospores in Trypetheliaceae. A1, Constrictolumina malaccitula; A2, C. majuscula, with ornamented wall; A3, Pseudopyrenula diluta, in part with yellow content in lumina; B, Bogoriella decipiens; C, Distothelia angulata, with thick terminal walls; D, Polymeridium pleurothecium, thin-walled; E, Trypethelium subeluteriae, thin-walled; F, Bathelium nigroporum, thin-walled; G, Astrothelium diplocarpoides, with diamond-shaped lumina; H, Architrypethelium nitens, with crystal-like structures; I, Astrothelium megaspermum, muriform with rounded to diamond-shaped lumina; J & K, Aptrootia terricola, muriform, brown; in J young, still hyaline ascospore showing astrothelioid stage; L, A. robusta, muriform, brown; outer wall breaking apart. Scales: A–F=10 µm; G–L=20 µm. (Images A & C by F. Schumm).

Most species in Trypetheliaceae have colourless ascospores, but brown ascospores are found in the basal genera Bogoriella (Fig. 9B) and Novomicrothelia, in most species of Architrypethelium, and in Aptrootia (Fig. 9K & L), in which the large, muriform ascospores also form a peculiar, easily breakable outer shell (Fig. 9L) (Sweetwood et al. 2012).

Chemistry

The chemistry of the Trypetheliaceae is rather simple compared to other tropical, crustose families, such as Graphidaceae (Rivas Plata et al. 2012; Lumbsch et al. 2014), and secondary substances are restricted to two main groups: xanthones and pigments, mostly anthraquinones.

Lichexanthone is the most commonly found xanthone in the family. It reacts UV+ yellow and can be present on the ascomata, pseudostromata and/or the ostioles, and/or on the thallus (Fig. 10A), rarely in the medulla. The taxonomic value of the presence of lichexanthone has been disputed; species were traditionally separated based on this feature, in others it was considered infraspecific variation (e.g. Harris 1995). Based on evidence from phylogenetic data (Nelsen et al. 2014: Lücking et al. 2016 a ), we consider the presence and location of lichexanthone species-specific. 1,8-dihydroxy-3,6-dimethoxyxanthone (coronatone) is less often present; it reacts UV+ orange but might be masked by the present of lichexanthone. The presence and proportion of both substances in UV+ taxa needs to be studied further.

Fig. 10 Secondary chemistry and pigments in Trypetheliaceae. A, lichexanthone, Astrothelium phlyctaena. B–G, yellow-orange to red anthraquinones on ascomata and pseudostromata; B, A. croceum; C, A. kunzei; D, Marcelaria cumingii; E, M. purpurina; F, Astrothelium aurantiacocinereum; G, Trypethelium eluteriae. H, pseudostromata with yellow internal and orange and red external anthraquinone pigments, T. astroideum; I, fused ostiole with red quinone isohypocrellin, Astrothelium purpurascens; J, ascomata with pockets of red pigment, A. sierraleonense. K & L, pseudostromata with internal yellow and red pigment; K, A. degenerans; L, A. sanguinarium with isohypocrellin. Scales=1 mm.

Various quinones, mostly anthraquinones, are regularly present on or in the ascomata or pseudostromata, and/or the ostioles and/or on or in the thallus (Fig. 10B–L), sometimes even in the ascoma wall, the hamathecium, or inside the ascospores (Fig. 9A). Many anthraquinones have not yet been identified, especially when quantities are low (e.g. when restricted to the ostiole or ascospore). However, the following substances have been classified (Mathey & Hoder 1978; Mathey 1979; Mathey et al. 1980):

Parietin (=physcione) is the most common yellow-orange pigment, for example in Astrothelium aeneum, A. croceum (Fig. 10B), Marcelaria cumingii (Fig. 10D), and Trypethelium eluteriae (Fig. 10G); it reacts K+ purple.

Teloschistin (=fallacinol) is also yellow and is found in Marcelaria benguelensis; it reacts K+ purple.

Xanthorin (=lauropurpurone) is red and reacts K+ purple; it is found, for example, in Marcelaria purpurina (Fig. 10E).

Secalonic acid derivates are yellow to orange and react K+ yellow; they occur in Marcelaria purpurina.

Emodin and derivates are orange and react K+ purple; these pigments are found, for example, in Marcelaria cumingii (Fig. 10D).

The perylene quinone isohypocrellin is red and reacts with a K+ green efflux: this pigment occurs in Astrothelium purpurascens (Fig. 10I), A. sanguinarium (Fig. 10L), A. sanguineoxanthum, and Dictyomeridium isohypocrellinum.

Skyrin, isopigmentosin A and C, and semivioxanthin are also occasionally found; since these substances are detected in low concentration, they have not yet been assigned taxonomic value, since their absence or presence on chromatographic plates might depend on their concentration in the lichen and on analytical conditions. However, it is expected that detailed chemical analyses with sophisticated methods could reveal characteristic, species-specific patterns including these substances.

With regard to UV testing, it should be stressed that only positive UV-reactions that relate to xanthones and anthaquinones are mentioned in the descriptions. These are yellow or orange for xanthones and usually red for pigments; whitish or greenish reflections of the thallus or ascoma surface are ignored as these correspond to brightening under the UV lamp of ochraceous (often old) specimens; taxa with such UV reflections only are reported as UV−. Similarly, some specimens with a thick, hyaline cortex show a faint, K+ yellow reaction, which is also ignored as it seems to be caused by a structural change from wetting, not by a secondary substance. When carrying out UV testing, it is therefore recommended to make comparisons with previously identified reference specimens of taxa with a set of known chemical compounds, including lichexanthone and the major pigments.

Species Delimitation and Nomenclature

Species delimitation within Trypetheliaceae has been rather inconsistent in the past and, until recently, sometimes used the same characters to either separate species or include specimens within a single taxon (Harris 1993, 1995). This was also reflected at the genus level, where species with lateral ostioles and transversely septate ascospores were assigned to a single genus, Astrothelium, whereas those with lateral ostioles and muriform ascospores were assigned to two separate genera, Campylothelium (with solitary ascomata) and Cryptothelium (with fused ascomata).

To obtain a more consistent species concept we used two approaches: 1) analysis of morphological, anatomical and chemical variation in phylogenetically defined clades (Lücking et al. 2016 a ); and 2) morphological analysis of a larger number of specimens in larger species complexes. As a result, we found that characters such as thallus and ascoma morphology (e.g. surface structure and emergence), hymenial inspersion, ascospore size, and secondary chemistry (including lichexanthone) are diagnostic at the species level, much more so than previously believed; hence, a much larger number of species is recognized here based on a revised species concept alone (exluding the many novel taxa described elsewhere in this issue). For example, the rich material available from Rondônia and Sergipe in Brazil (Aptroot & Cáceres 2016; Aptroot et al. 2016 b ), with several hundred collections representing over 70 taxa, and from Venezuela (Komposch & Hafellner 1999; Komposch et al. 2002), with c. 300 collections belonging to 40 species, allowed us to analyze morphological and anatomical variation in some of the common taxa. As an example, the ascospore width and length of all specimens of the Astrothelium conicum-aggregate were analyzed showing that the species recognized within this complex are separated on a combination of UV-reaction and ascospore dimensions.

To obtain a stable nomenclature that reflects this revised species concept, as many collections as possible were studied including most type specimens of published names. Fifty-seven new lectotypes were designated, including in one case an illustration with an additional epitype. To check possibly erroneous nomenclatural citations, including those given in Index Fungorum and other similar databases, original protologues of all names were studied. As an extreme example of a revised species concept and nomenclatural treatment, Astrothelium variolosum sensu Harris (1993), which in most other treatments corresponds to the four names Astrothelium confusum, A. variolosum (ostioles lateral, fused, without or with lichexanthone), Trypethelium nitidiusculum and T. ochroleucum (ostioles apical, separate, without or with lichexanthone), is now considered to encompass at least 34 different species (Table 1) which are not necessarily closely related (Lücking et al. 2016 a ). The fact that all of these names were described in the 19th century underlines the importance of critical analysis of historical literature and type specimens to accurately assess the taxonomy and nomenclature of Trypetheliaceae. This situation is similar to that found in the genus Ocellularia where the commonly used names O. papillata, O. perforata and O. terebrata now refer to 69 different taxa, many of them represented by historical names (Lücking 2014).

Table 1 Revised species concept in the Trypethelium nitidiusculum/T. ochroleucum-Astrothelium variolosum complex

*Lichexanthone location: +/+ indicates lichexanthone present on both thallus and ascomata (pseudostromata); +/– indicates lichexanthone present on the thallus but not on the ascomata (pseudostromata) and –/+ indicates lichexanthone not present on the thallus but is present on the ascomata (pseudostromata).

Keys and short descriptions are presented for the currently accepted species of Trypetheliaceae with the exception of those described elsewhere in this issue as new taxa; these are only cross-referenced in the keys. The delimitation of Astrothelium, Bathelium, and Trypethelium is revised so as to be in concordance with the molecular phylogeny, resulting in mostly monophyletic genera that are still morphologically recognizable. Many taxa, including the genera Campylothelium, Cryptothelium and Laurera, are synonymized for the first time (the latter three with Astrothelium) whereas others, including the genus Bogoriella, have been reinstated.

The revised genus and species concept and the examination of many old type specimens led to the proposal of many new combinations. Numerous species are recorded for the first time from a country or continent. However, there is still a high proportion of species known only from their type material. This also strongly suggests that many species still remain undiscovered, as shown by a statistical prediction exercise (Aptroot et al. 2016 c ). This is corroborated by the fact that a few short field trips in small areas in Brazil yielded many undescribed species (Aptroot & Cáceres 2016; Aptroot et al. 2016 b ; Lücking et al. 2016 a ), and also taxonomic inventories of other understudied areas, such as Panama and Bolivia, unravelled numerous novel taxa (Flakus et al. 2016; Lücking et al. 2016 b ).

Some commonly used names are synonymized for the first time, such as Trypethelium ochroleucum; many rarely applied names are reinstated, as in the example of the Astrothelium variolosum complex above. Inevitably this resulted in new names for some of the most common taxa. We preferred this solution, over the alternative of proposing well-known names for conservation, for two reasons: 1) Trypetheliaceae are relatively rarely mentioned in the general literature and mostly known to specialists; 2) the new names reflect the revised taxonomy and hence will force future workers to assess morphological, anatomical and chemical characters more critically. For instance, the name Trypethelium ochroleucum has been applied in a pantropical context to a number of species now recognized as different: Astrothelium porosum (pseudostromatic, inspersed), A. pulcherrimum (solitary, clear) and A. phlyctaena (pseudostromatic, clear). The fact that the name ochroleucum has been subsumed in synonymy will hopefully lead to proper revision and identification of herbarium material and newly collected specimens. Similarly, the taxon formerly known as Trypethelium nitidiusculum was split into Astrothelium bicolor (pseudostromatic, clear), A. nitidiusculum (solitary, clear) and A. scoria (grouped, inspersed).

We are very much aware that the present revision is not a thorough monograph of Trypetheliaceae. Rather, we present this revisionary synopsis to stimulate further research in this highly diverse and fascinating family, hoping that future workers will pay more attention to morphological, anatomical and chemical features to identify species and that the molecular phylogenetic data will be much expanded. Undoubtedly, further research will lead to additional changes in the classification of Trypetheliaceae and many additional new species will be discovered (Aptroot et al. 2016 c ). Notably, we are more confident with the taxonomic concept of the less common and rare species, which often have a unique combination of diagnostic characters different from similar and related taxa, than with the common species, such as Astrothelium aeneum, A. bicolor, A. porosum, A. phlyctaena, A. scoria and Trypethelium eluteriae which have a degree of variation that needs to be assessed by additional molecular phylogenetic studies.

Distribution and Ecology

Trypetheliaceae is almost exclusively tropical and epiphytic, with very few species (e.g. Viridothelium virens) found in temperate regions and few species growing on other substrata. In general terms, the distribution and ecology of Trypetheliaceae is very similar to that of Graphidaceae although the latter has more taxa in extra-tropical regions and on substrata other than bark (Rivas Plata et al. 2008; Lücking et al. 2014). More commonly than Graphidaceae, Trypetheliaceae are found in (semi-) exposed microhabitats and habitats, such as the forest canopy and open savannahs and dry forest, often forming colourful crustose lichen communities with dominant taxa producing yellow to orange pigments (Komposch & Hafellner 2000, 2002, 2003; Cáceres 2007; Aptroot et al. 2008; Cáceres et al. 2008; Rivas Plata et al. 2008). It appears that species with green thalli and partially immersed or covered ascomata are more frequently found in the rainforest understorey, but this has not been tested quantitatively. However, the observation that many taxa are typical of the more exposed canopy, a microhabitat that has not been well studied, indicates that this microhabitat might harbour many yet unknown species.

Following this revisionary synopsis, the list of accepted species known from the various countries has changed considerably, especially for countries in which much previous work was carried out, such as India and Brazil. A table with the currently accepted species and their updated, confirmed world distribution is available as supplementary material in Aptroot et al. (2016 c ).

Some species are much more widespread and common than others. In order to facilitate identification, lists are given here of the most common species. These are only a fraction of those known. Half of what remains are known only from the type location; the chance of finding any of these elsewhere is probably not much higher than that of finding an undescribed species, as demonstrated by the papers on Bolivian and Rondônian Trypetheliaceae in this issue (Aptroot & Cáceres 2016; Flakus et al. 2016).

Species with over 100 specimens seen include: Astrothelium macrocarpum, A. phlyctaena, A. scoria, A. scorioides, A. versicolor, Constrictolumina cinchonae, Nigrovothelium tropicum and Trypethelium eluteriae.

Species with between 20 and 100 specimens seen include: Astrothelium aeneum, A. bicolor, A. cinnamomeum, A. degenerans, A. eustomum, A. feei, A. inspersaeneum, A. megaspermum, A. obscurum, A. ochrothelium, A. porosum, A. variolosum, Bathelium madreporiforme, Bogoriella hemisphaerica, B. punctata, B. subfallens, Dictyomeridium proponens, Marcelaria benguelensis, M. purpurina, Polymeridium albocinereum, P. catapastum, P. pleiomerellum, P. subcinereum, Pseudopyrenula diluta, P. subgregaria, P. subnudata, Trypethelium subeluteriae and Viridothelium virens.

Taxonomic Treatment

Trypetheliales Lücking, Aptroot & Sipman

In Aptroot et al., Biblioth. Lichenol. 97: 13 (2008); type: Trypetheliaceae Eschw.

Ascomycetes, usually lichenized with trentepohlioid algae, more rarely bark saprobes or lichenicolous fungi.

Ascomata perithecia, solitary or grouped and fused (with separate or fused ostioles), round, immersed to sessile. Wall dark brown to carbonized. Interascal hyphae thin, richly branched and anastomosing, forming a net-like structure. Asci fissitunicate, I−. Ascospores ellipsoid-fusiform, with variously developed endospore thickenings giving the lumina a diamond-shaped or more rarely a round outline, colourless or rarely (dark) brown.

Discussion. This order encompasses a phylogenetically quite distinct group, now containing two families, viz. the lichenicolous Polycoccaceae and the lichenized Trypetheliaceae (Aptroot et al. 2008; Hyde et al. 2013; Ertz et al. 2015).

The order Trypetheliales, based on Trypetheliaceae Eschw., was introduced by Aptroot et al. (2008) to reflect the phylogenetic distinctiveness of the Trypetheliaceae within Dothideomycetes (Del Prado et al. 2006; Nelsen et al. 2009, 2011). The name Trypetheliaceae Eschw. itself (Eschweiler 1824) has been considered invalidly published in Index Fungorum (IF 81884) and MycoBank (MB 81884), with the argument that it was associated with the rank term “cohors”; the same applies to the family names Parmeliaceae Eschw., Usneaceae Eschw., and Verrucariaceae Eschw. However, this interpretation appears to be incorrect, since an inappropriate rank term does not invalidate a name. Rather, ICN Art. 17.2 and 18.2 specify that if there is conflict between the intended rank (based on context and/or Latin termination) and the associated rank term, a name is presumed to be validly published at the intended rank, but not invalid. According to Art. 17.2, one might then conclude that Trypetheliaceae Eschw. (Eschweiler 1824) is to be accepted as a valid name at the order level (i.e. Trypetheliales Eschw.) because of the associated rank term “cohors”. In that case, because the principles of priority and homonymy do not apply to names above the rank of family, the names Trypetheliales Lücking et al. and Trypetheliales Eschw. would both be legitimate and could be used interchangeably. However, the opposite is the case, since Art. 17.2 states: “Names intended as names of orders, but published with their rank denoted by a term such as ‘cohors’, ‘nixus’, ‘alliance’, or ‘Reihe’ instead of ‘order’, are treated as having been published as names of orders.” This means that a name is to be treated at the order level, if the intention was to designate a name as an order, in spite (not because) of having used a rank term such as “cohors” (i.e. ‘cohors’ is not a rank term that automatically designates the rank of order). There is no further definition of the term ‘cohors’ in the Code, but its best Latin translation would be ‘group’, and there is no provision in the Code that the use of the term ‘cohors’ would preclude the meaning of family (as a group of genera). Correspondingly, the correct interpretation of the analogous Art. 18.2 is that a name is to be treated as family if the intention was to designate a family, in spite of using the rank term ‘ordo’. Eschweiler’s (1824) intention undoubtedly was to establish a family, as he used the term “familia” in the Latin description, defined Trypetheliaceae as a group of genera (the next higher, principal hierarchical level above genus is family), and used the correct Latin termination “-aceae” (Art. 18.1); technically, the termination is not relevant as the Code was not in place at the time, but the termination ‘-aceae’ has been widely accepted since the late 18th century as denoting taxa at the family level in botanical nomenclature. As a consequence, neither Art. 17.2 nor Art. 18.2 apply in this case, since Trypetheliaceae Eschw. was not intended as a name of an order (required for Art. 17.2) and it was not associated with the rank term ‘ordo’ (required for Art. 18.2). On the other hand, by extension of the meaning of both articles, which imply that 1) ‘cohors’ is not a rank term naturally corresponding to order level, 2) a name is to be accepted at the level of family if intended so, and 3) there is no conflict between the use of the rank term ‘cohors’ and the family rank defined in either article or the Code as a whole, the name Trypetheliaceae Eschw. (Eschweiler 1824) is valid at the family rank and is not to be treated at the order level or to be considered invalid. Similar considerations would apply to the names Parmeliaceae Eschw., Usneaceae Eschw., and Verrucariaceae Eschw.

Trypetheliaceae Eschw.

Eschweiler, Syst. Lich.: 17 (1824); Fée, Essai Crypt. Écorc.: xxvi (1824; as “Trypetheliées”), nom. inval. [ICN Art. 18.4, 32.1(b)]; Trypetheliaceae Zenker, in Goebel & Kunze, Pharmaceutische Waarenkunde 1(3): 123 (1827; as “Trypethelia”). nom. illeg. [ICN Art. 52.1]; type: Trypethelium Spreng.

Thallus crustose, corticate or not, without pseudocyphellae, sometimes surrounded by a black hypothallus. Algae trentepohlioid.

Ascomata perithecioid, mostly simple, but sometimes with fused ostioles, or the ascomata fused and having shared ostioles, with or without pseudostromatic tissues of a different structure and colour. Ostioles apical or eccentric. Wall usually completely carbonized, rarely only partly carbonized, with or usually without a clypeus, with or without crystals. Hamathecium usually colourless, rarely yellow, IKI−, clear or inspersed with oil droplets. Hamathecial filaments branched and usually anastomosing paraphysoids of 0·5–2·0 μm thick, rarely almost unbranched paraphysoids that are much wider at the base than at the tips; periphyses absent; short periphysoids rarely present. Asci bitunicate, cylindrical, IKI−, usually with a broad ocular chamber that is especially distinct in immature asci. Ascospores 1–8 per ascus, distoseptate, usually IKI−, rarely IKI+ violet, with rounded or angular lumina, without additional eusepta, colourless, rarely becoming brown, rarely constricted at the septa, with or without a gelatinous sheath.

Conidiomata pycnidia, containing microconidia. Conidia colourless, rod-shaped. Conidiogenesis acrogenous.

Chemistry. Lichexanthone (or the closely related 1,8-dihydroxy-3,6-dimethoxyxanthone or coronatone) or quinones, mostly anthraquinones, often present. Non-specific substances such as skyrin, isopigmentosin A and C and semivioxanthin sometimes detected.

Discussion. This family currently comprises 16 genera. Most were traditionally assigned to this family, but some have been erected here to accommodate species aggregates traditionally assigned to other groups (e.g. Arthopyrenia, Mycomicrothelia) that were found to belong here based on phylogenetic studies (Nelsen et al. 2009, 2011, 2014; Lücking et al. 2016 a ). The new genus Constrictolumina is used for species formerly placed in Arthopyrenia, which in its strict sense remains a small, non-lichenized, extratropical genus (Hyde et al. 2013), and Novomicrothelia for several species previously classified in Mycomicrothelia, while Bogoriella is reinstated to accommodate most of the former Mycomicrothelia species. Most genera accepted here are supported by molecular data, whereas the inclusion of the reinstated Distothelia is based on morphological similarities. The circumscription of Astrothelium, Bathelium, and Trypethelium had to be changed so as to be in concordance with the phylogeny (Nelsen et al. 2014; Lücking et al. 2016 a ). Bathelium and Trypethelium are restricted here to small, morphologically recognizable groups of species related to their respective types; some species included in Bathelium sensu Harris (1995), with astrothelioid ascospores, were found to belong in Astrothelium (Nelsen et al. 2014; Lücking et al. 2016 a ). The new genus Viridothelium encompasses a clade with species morphologically similar to Astrothelium but with ascospores reminiscent of those of Trypethelium, whereas Marcelaria was recently established for a small group of species with very conspicuous ascomata and muriform ascospores (Aptroot et al. 2013 a ).

For the correct nomenclature and authorship of the name Trypetheliaceae, see discussion under Trypetheliales above.

Aptrootia Lücking & Sipman

In Lücking et al., Lichenologist 39: 188 (2007); type: Aptrootia terricola (Aptroot) Lücking, Umaña & Chaves (holotype).

Thallus either corticate and green and bullate or verrucose, or not corticate and grey, ±smooth and cartilaginous.

Ascomata solitary, black, globose to ampulliform, immersed in thalline warts or in the substratum. Ostioles apical. Hamathecium colourless, clear or inspersed with large irregular oil droplets near the ostiole, filaments thin, anastomosing paraphysoids. Ascospores 1(–2) per ascus, IKI+ violet, with scarcely rounded lumina, initially colourless, becoming dark brown but wall internally remaining hyaline (only outer layer brown), ornamented with brown warts or not, fusiform or usually elongate-ellipsoidal to bacilliform with subacute or rounded ends, densely irregularly muriform, not constricted at the septa, surrounded by a gelatinous sheath.

Pycnidia unknown.

Chemistry. No substances detected.

Discussion. This recently established genus (Lücking et al. 2007) differs from other Trypetheliaceae in the combination of immersed ascomata and dark brown, muriform ascospores, as well as in its peculiar ecology growing often over bryophytes, a feature otherwise unknown in the family. The genus was erected for a terricolous lichen known from mountains in Papua New Guinea and Costa Rica. Two additional Australasian species have been transferred to Aptrootia (Aptroot 2009 a ).

Aptrootia elatior (Stirt.) Aptroot

Fl. Australia 57: 600 (2009).—Ascidium elatius Stirt., J. Linn. Soc., Bot. 14: 466 (Feb. 1875).—Leptotrema elatius (Stirt.) Müll. Arg., Bull. Herb. Boissier 2 (App. 1): 75 (1894).—Thelotrema elatius (Stirt.) Hellb., K. Svenska Vetensk.-Akad. Handl. 21(3, 13): 79 (1896).—Laurera elatior (Stirt.) D. J. Galloway, New Zealand J. Bot. 21: 193 (1983); type: New Zealand, near Wellington, Buchanan (BM!—lectotype; Galloway, Fl. New Zealand Lichens: 205, 1985).

Ascidium melanosporum C. Knight, Trans. Proc. N. Z. Inst. 7: 363 (Jul. 1875); type: New Zealand, Knight s. n. (NSW!).

Anthracothecium monosporum Müll. Arg., Bull. Herb. Boissier 3: 327 (1895).—Polyblastiopsis monosporum (Müll. Arg.) Upreti & A. Singh, Brunonia 10: 226 (1987).—Julella monospora (Müll. Arg.) D. D. Awasthi, Lichenology in Indian Continent: 15 (2000); type: Australia, Victoria, Knight 214 (G!—holotype).

Verrucaria luteonitens Nyl., Ann. Soc. Sci. Fenn. 26(10): 24 (1900).—Anthracothecium luteonitens (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 464 (1922); type: Sri Lanka, Pedretalagalla, Almquist (H-Nyl 1061!—holotype).

(Fig. 11A)

Fig. 11 Habitus, anatomy and ascospore of Aptrootia (A–C) and Architrypethelium species (D–L). A, Aptrootia elatior (New Zealand, lectotype); B, A. terricola (Papua New Guinea, Aptroot 37658); C, A. robusta (Tasmania, Lumbsch 20012n); D, Architrypethelium grande (Brazil, lectotype); E, A. lauropaluanum (Brazil, holotype); F & G, A. hyalinum (Venezuela, Sipman & van der Werff 10902); H , A. penuriaxanthum (Bolivia, holotype); I & J, A. nitens (I, Venezuela, holotype of Pleurothelium ernstianum; J, Costa Rica, Lücking 15212b); K, A. columbianum (Colombia, lectotype); L, A. uberinum (Costa Rica, Lücking s. n.). Scales: A, B, D–F, H, I, K & L=1 mm; C=100 µm; G & J=50 µm.

Thallus corticolous, corticate, pale yellow-brown to green, verrucose-bullate.

Ascomata trypethelioid, with apical ostioles, ampulliform, oval or subglobose (in section), 0·7–1·7 mm diam., erumpent, covered by thallus except the rather broad, black ostiolar area, with copious hyaline crystals and a conspicuous black ring around the ostiole free of crystals. Wall to 150 µm thick. Hamathecium clear except for large irregular oil droplets near the ostiole. Ascospores 1(–2) per ascus, 200–330×60–90 µm, oblong-ellipsoid, richly muriform, becoming dark brown and ornamented (verruculose), with ascospore wall distinctly bi-layered: outer layer dark brown, inner layer hyaline.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (previously reported from Australia and New Zealand, now also from Sri Lanka and Sabah).

Discussion. This species differs from the other two by the ornamented ascospores and bark as substratum. We suspected that it may represent an undescribed genus, but it is phylogenetically nested between the other two species. The ascospore surface with the irregular brown warts is unique in the family. In a thin hand-section, this outer wall appears to be brittle, and a section through the ascospores reveals the subhyaline internal parts.

New country record. Malaysia: Sabah, Kota Belud, Kinabalu Park, 2800 m, 1989, Sipman & Tan 31305 (B).

Aptrootia robusta (P. M. McCarthy & Kantvilas) Aptroot

Fl. Australia 57: 661 (2009).—Laurera robusta P. M. McCarthy & Kantvilas, Lichenologist 25: 51 (1993); type: Australia, Tasmania, Crater Peak, Kantvilas & James (HO—holotype, not seen; BM!—isotype).

(Fig. 11C)

Thallus muscicolous, with thin, cartilaginous cortex, grey, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, globose (in section), 0·9–1·5 mm diam., immersed, partially covered by thallus except the rather broad, black ostiolar area. Wall to 120 µm thick. Hamathecium clear. Ascospores 1 per ascus, 150–360(–400)×65–140 µm, ellipsoid, richly muriform, becoming dark brown.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Australia.

Discussion. This species differs from Aptrootia terricola principally by the larger ascospores (170–230×40–70 µm in the latter).

Aptrootia terricola (Aptroot) Lücking et al.

In Lücking et al., Lichenologist 39: 188 (2007).—Thelenella terricola Aptroot, Fungal Diversity 2: 45 (1999); type: Papua New Guinea, Simbu Prov., Mount Wilhelm, Pindaunde Valley, near the hut on the S-shore of Lake Piunde, Aptroot 32649 (CBS!—holotype; ABL!—isotype).

(Fig. 11B)

Thallus terricolous, with thin, cartilaginous cortex, grey, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, globose (in section), 0·9–1·5 mm diam., immersed, partially covered by thallus except the rather broad, black ostiolar area. Wall to 120 µm thick. Hamathecium clear. Ascospores 1 per ascus, 170–230×40–70 µm, ellipsoid, richly muriform, becoming dark brown late during maturity, I+ violet when hyaline.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Amphi-Pacific, known from tropical mountains in Costa Rica and Papua New Guinea, now also reported from the Solomon Islands.

Discussion. This species differs from Aptrootia robusta principally by the smaller ascospores (150–360(–400)×65–140 µm in the latter).

New country record. Solomon Islands: Guadalcanal Island: Central part, Mount Popomansiu, summit, c. 2200 m, 1965, Hill 9400 (BM, ABL).

Architrypethelium Aptroot

Biblioth. Lichenol. 44: 120 (1991); type: Architrypethelium uberinum (Fée) Aptroot (holotype).

Thallus corticate.

Ascomata solitary or aggregated. Ostioles apical or eccentric. Wall hyphal (textura intricata), carbonized. Hamathecium clear or inspersed with oil droplets, filaments thin, anastomosing paraphysoids. Ascospores distoseptate, usually also with euseptate walls, mostly brown, 3–5-septate, large, often with longitudinal folds in the wall, rarely colourless.

Pycnidia unknown.

Chemistry. Lichexanthone rarely present.

Discussion. Architrypethelium externally resembles species of Astrothelium, including those previously placed in the genera Laurera, Cryptothelium, and Trypethelium, but differs anatomically by its 3-septate, extremely large ascospores, which are hyaline to dark brown when mature and do not have the diamond-shaped lumina typical of Astrothelium when mature (Aptroot 1991). Phylogenetically it can be considered the sister clade to Astrothelium. Species of Pyrenula with large, 3-septate ascospores can be confused with Architrypethelium but, in addition to the differences in hamathecium structure, are distinguished as follows: Pyrenula subpraelucida has ascospores with small terminal lumina against the endospore, while in P. laii and P. montocensis, the ascospores have angular lumina with very thick septa and lateral walls. In addition, 3-septate ascospores in Pyrenula are less than 90 µm long while in Architrypethelium they are usually longer than 90 µm (up to 160 µm).

Architrypethelium columbianum (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816650

Trypethelium columbianum Nyl., Ann. Sci. Nat. Bot. sér. 5, 3: 347 (1867); type: Colombia, Rio Negro, Lindig 35 (H-Nyl 321!—lectotype, designated here; BM!, BR—isolectotypes).

(Fig. 11K)

Thallus corticate, light olive-grey, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·7–1·2 mm diam., sessile, covered by a brownish thallus layer. Wall thick, fully carbonized. Hamathecium inspersed. Ascospores 2–4 per ascus, 120–155×35–50 µm, oblong-ellipsoid, 3-septate, outer lumina much smaller than inner lumina, lumina rounded in the corners, often with needle-like crystals, wall 3–5 µm thick, surrounded by a gelatinous sheath 7–10 µm wide, becoming dark brown, I−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Colombia).

Discussion. This species differs from Architrypethelium uberinum mainly in the inspersed hamathecium.

Architrypethelium grande (Kremp.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816651

Ascidium grande Kremp., Flora 59: 249 (1876); Phaeotrema grande (Kremp.) Zahlbr., Catal. Lich. Univ. 2: 607 (1923); type: Brazil, Rio de Janeiro, Glaziou 6271 (M!—lectotype, designated here; BM!, C!—isolectotypes).

(Fig. 11D)

Thallus corticate, olive-green to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 1–3 mm diam., sessile, covered by a light, orange-brown layer. Wall thick, fully carbonized, with thick, gelatinous cortex composed of anastomosing hyphae. Hamathecium clear. Ascospores 2–4 per ascus, 120–165×40–50 µm, oblong-ellipsoid, 3-septate, lumina angular to almost rounded in the corners, often with needle-like crystals, wall 3–5 µm thick, surrounded by a gelatinous sheath 3–5 µm wide, hyaline (although given as becoming pale olive-brown in the protologue), I−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Discussion. This species differs from the other two species with hyaline ascospores, Architrypethelium hyalinum and A. penuriaxanthum, in the sessile ascomata, and from A. hyalinum also in the absence of lichexanthone. Krempelhuber (1876) described the ascospores as becoming pale olive-brown, but we observed only hyaline ascospores in the type material. The only species with brown ascospores, an apical ostiole and clear hymenium is A. uberinum, from which A. grande also differs markedly in the sessile, orange-brown perithecial warts.

Architrypethelium hyalinum Aptroot

In Aptroot et al., Biblioth. Lichenol. 98: 37 (2008); type: Costa Rica, Puntarenas, Las Cruces Biological Station, trail to Rio Java, Sipman 53229 (B!—holotype; INB-3993295!—isotype).

(Fig. 11F & G)

Thallus corticate, olive-green, smooth to uneven, sometimes gall-like.

Ascomata trypethelioid, with apical ostioles, solitary, 0·7–1·5 mm diam., prominent, irregularly covered by thallus except for the black ostioles which are sometimes surrounded by an ochraceous zone. Wall 200–400 µm thick, fully carbonized. Hamathecium clear. Ascospores 4–8 per ascus, 100–150×30–50 µm, oblong-ellipsoid, often curved, 3-septate, outer lumina much smaller than inner lumina, lumina rounded in the corners, often with needle-like crystals, constricted at the median septum, wall 3–5 µm thick, surrounded by a gelatinous sheath 7–15 µm wide, hyaline, I−.

Chemistry. Thallus and ascomata UV+ yellow, K−. TLC: lichexanthone.

Distribution. Neotropical (previously reported from Costa Rica and Brazil).

Discussion. This Architrypethelium has among the largest 3-septate ascospores in the family. It is also the only species in the genus with lichexanthone. The species could be mistaken for a Trypethelium except for the extremely large ascospores.

New country records. Puerto Rico: Distr. Mayagüez: Reserva Forestal Maricao, 1989, Aptroot & Aptroot 24960, 25322 (ABL).—Venezuela: Falcon: Miranda, 1979, Sipman & van der Werff 10902 (B).—Colombia: Santuario, 1989, Wolf et al. 5992 (ABL, COL).—Guyana: Demerara-Berbice District: Mabura Hill, 1988, Bleij & Biesmeijer s. n. (ABL).

Architrypethelium nitens (Fée) Aptroot

In Aptroot et al., Biblioth. Lichenol. 98: 38 (2008).—Verrucaria nitens Fée, Essai Crypt. Écorc.: 88 (1824); type: South America, on “cinchona” (L!—lectotype; Aptroot et al., Biblioth. Lichenol. 98: 38 (2008)).

Pyrenastrum seminudum Mont., Ann. Sci. Nat. Bot. sér. 2, 19: 64 (1843).—Architrypethelium seminudum (Montagne) Aptroot, Biblioth. Lichenol. 44: 120 (1991); type: French Guiana, Leprieur 588 (PC!—holotype).

Pleurothelium ernstianum Müll. Arg., Flora 60: 475 (1877).—Parathelium ernstianum (Müll. Arg.) Müll. Arg., Hedwigia 34: 36 (1895); type: Venezuela, Caracas, Ernst 130 (G!—holotype).

Parathelium superans Müll. Arg., Bull. Soc. Roy. Bot. Belg. 32(1): 165 (1893).—Splanchnonema superans (Müll. Arg.) O. Erikss., Opera Bot. 60: 133 (1981); type: Costa Rica, Puntarenas, Boruca, Tonduz s. n. (G!—lectotype, designated here; BR!, G!—isolectotypes; Pittier, Pl. Costaric. Exs. 6253).

(Fig. 11I & J)

Thallus corticate, olive-green, smooth to uneven.

Ascomata pleuro- to astrothelioid, solitary with eccentric ostioles or few chambers joined with eccentric, fused ostioles, 1·0–1·5 mm diam., erumpent to prominent, covered by thallus except for the blackish ostiolar area. Wall thick, fully carbonized. Hamathecium clear. Ascospores 8 per ascus, 90–150×25–50 µm, oblong-ellipsoid, 3-septate, outer lumina much smaller than inner lumina, lumina rounded in the corners, often with needle-like crystals, wall 3–5 µm thick, surrounded by a gelatinous sheath 7–10 µm wide, becoming dark brown, I−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Costa Rica, Venezuela, Guyana, French Guiana and Brazil).

New country records. Puerto Rico: Mayagüez: Reserva Forestal Maricao, 1989, Aptroot & Aptroot 24930, 25420, 25040, 25002 (ABL).—Panamá: Panamá: Alto de Campana, 2001, Etayo 18782 (ABL).

Architrypethelium uberinum (Fée) Aptroot

Biblioth. Lichenol. 44: 122 (1991).—Porina uberina Fée, Essai Crypt. Écorc.: 83 (1824).—Porophora uberina (Fée) Spreng., Syst. Orb. Veget. 4(1): 242 (1827).—Pyrenula uberina (Fée) Fée, Essai Crypt. Écorc. Suppl.: 84 (1837).—Pertusaria uberina (Fée) A. Massal., Ric. Auton. Lich. Crost.: 190 (1852).—Trypethelium uberinum (Fée) Nyl., Mém. Soc. Sci. Nat. Cherbourg 5: 141 (1857).—Verrucaria uberina (Fée) Trevis., Consp. Verruc.: 8 (1860).—Stromatothelium uberinum (Fée) Trevis., Flora 44: 20 (1861).—Pseudopyrenula uberina (Fée) Vain., Ann. Acad. Sci. Fenn., ser. A 15(6): 353 (1921); type: Peru, “cort. cinchonae” (G!—holotype).

Trypethelium uberinoides Nyl., Flora 41: 381 (1858); type: Mexico, Orizaba, Müller (PC—lectotype, designated here).

Verrucaria megalospora Kremp., Flora 59: 525 (1861).—Parathelium megalosporum (Kremp.) Müll. Arg., Hedwigia 32: 134 (1893); type: Brazil, Rio de Janeiro, Glaziou 6276 (M!—holotype; BM!, UPS!—isotypes).

(Fig. 11L)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, 1·0–1·7 mm diam., erumpent to prominent, covered by thallus except for the blackish ostiolar area surrounded by a whitish rim. Wall thick, fully carbonized. Hamathecium clear. Ascospores 2–4 per ascus, 120–155×35–50 µm, oblong-ellipsoid, 3-septate, outer lumina much smaller than inner lumina, lumina rounded in the corners, often with needle-like crystals, wall 3–5 µm thick, surrounded by a gelatinous sheath 7–10 µm wide, becoming dark brown, I−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Possibly pantropical; previously reported from Costa Rica, Peru, and Brazil. Incorrectly reported from Sri Lanka but now found in Oceania.

New country records. American Samoa: Tutuila: Pago Pago, 1970, Degelius P-279 (UPS).—French Polynesia: Tahiti: Taravao, 1970, Degelius P-371 (UPS).

Astrothelium Eschw.

Syst. Lich.: 18 (1824); type: Astrothelium conicum Eschw. [=A. cinnamomeum (Eschw.) Müll. Arg., lectotype; Massalongo, Atti I. R. Istitut. Veneto, Ser. 3, 5: 335, 1860].

Meissneria Fée, Essai Crypt. Écorc. Suppl.: 66 (1837), nom. illeg., non Meisneria DC. (1828); type: Trypethelium deforme Fée (as Meissneria varia Fée, holotype).

Laurera Reichenb., Deutsch. Bot. Herb.-Buch: 15 (1841), nom. cons., nom. nov. pro Meissneria Fée; type: Trypethelium deforme Fée (as Meissneria varia Fée, holotype).

Heufleria Trevis., Spighe e Paglie: 19 (1853); type: Heufleria conica (Eschw.) Trevis. (= cf. Astrothelium cinnamomeum, holotype).

Meristosporum A. Massal., Atti I. R. Istitut. Veneto, Ser. 3, 5: 327 (1860); type: Meristosporum javanicum A. Massal. (holotype).

Cryptothelium A. Massal., Atti I. R. Istitut. Veneto, Ser. 3, 5: 335 (1860); type: Cryptothelium sepultum (Mont.) A. Massal (holotype).

Leightonia Trevis., Flora 44: 19 (1861), nom. illeg., non. Trevis. (1853); type: Leightonia porosa (Ach.) Trevis. (holotype).

Campylothelium Müll. Arg., Flora 66: 245 (1883); type: Campylothelium puiggarii Müll. Arg. (holotype).

Thallus corticate, mostly olive-green.

Ascomata simple or aggregated or forming pseudostromata, immersed to prominent, with the pseudostromata often of a different structure and colour than the thallus. Ostioles apical or eccentric, simple or fused. Wall hyphal (textura intricata), usually carbonized. Hamathecium clear or inspersed with oil droplets, filaments thin, anastomosing paraphysoids. Ascospores distoseptate, hyaline, transversely septate or muriform.

Pycnidia occasionally present.

Chemistry. Lichexanthone, red crystals and/or anthraquinones regularly present, in or on the thallus, pseudostromata and/or ostioles.

Discussion. The type species, Astrothelium conicum, was not validly published when the genus Astrothelium was published (Eschweiler 1824), but only later by Eschweiler (in Martius 1833); however, this does not render the description of the genus invalid. Several other species were described in the genus prior to A. conicum, which makes the lectotypification with the latter slightly unusual, but not invalid either. The synonymy of A. conicum (the type species of the genus) with A. cinnamomeum was confirmed by combining the observations of Müller (1884: 270), who observed the ascospores, and Harris (1986: 59), who observed no ascospores but saw the negative reaction with UV.

Astrothelium acrophaeum Müll. Arg.

Bot. Jahrb. Syst. 6: 383 (1885).—Trypethelium acrophaeum Nyl. (1876), nom. nud.—Pleurotrema acrophaeum (Müll. Arg.) R. C. Harris, Acta Amazon. (Suppl.) 14: 69 (‘1984’) [1986]; type: Cuba, Wright s. n. (G!—holotype; H-Nyl 268!, US!—isotypes; Müller, Verr. Cub. 176).

(Fig. 28B)

Thallus corticate, light olive-green to yellowish, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to irregularly confluent or diffusely pseudostromatic, 0·7–1·1 mm diam., erumpent to prominent, basally to laterally covered by thallus but upper part and ostiolar area exposed and blackish. Hamathecium clear. Ascospores 8 per ascus, 2(–3)-septate, fusiform, 12–14×4–5 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (reported from Cuba).

Astrothelium aeneum (Eschw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816652

Verrucaria aenea Eschw., in Martius, Icon. Sel. Pl. Crypt. 2: 15, t. 4, fig. 3 (1828).—Pseudopyrenula aenea (Eschw.) Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 207 (1890).—Trypethelium aeneum (Eschw.) Zahlbr., in Engler & Prantl, Nat. Pflanzenfam. 1(1*): 70 (1903); type: Brazil, Bahia, Caetité, Martius s. n. (M!—holotype).

Verrucaria tetracerae var. crocea Eschw., in Martius, Flora Brasil. 1: 134 (1833).—Spermatodium croceum (Eschw.) Trevis., Conspect. Verruc.: 6 (1860); type: Brazil, Martius s. n. (M—holotype, not seen).

Verrucaria myriococca Spreng., Syst. Veget. 4(1): 245 (1827); type: not found in L, probably lost.

Verrucaria heterochroa Mont., Ann. Sci. Nat., Bot., sér. 2, 19: 60 (1843).—Pyrenula heterochroa (Mont.) Trevis., Spighe e Paglie: 17 (1853).—Segestria heterochroa (Mont.) Trevis., Conspect. Verruc.: 6 (1860).—Pseudopyrenula heterochroa (Mont.) Mull. Arg., Flora 66: 248 (1883); type: French Guiana, Leprieur 452 (BM—lectotype, designated here).

(Fig. 15B–F)

Thallus corticate, light olive-green to yellowish but largely covered with orange pigment, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent or diffusely pseudostromatic, 0·3–0·7 mm diam., erumpent, covered by thallus and orange pigment except for dark ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 20–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ red, K+ purple, with anthraquinone.

Distribution. Pantropical (previously reported from the USA, Costa Rica, El Salvador, Islas Revillagigedo, Bahamas, Cuba, Jamaica, Colombia, Venezuela, Guyana, Surinam, French Guiana, Galapagos, Bolivia, Brazil, Sri Lanka, Thailand, Malaysia (Sarawak, Sabah), Papua New Guinea, and Australia).

Discussion. Astrothelium aeneum as defined here might still represent a collective taxon, including forms with solitary or almost pseudostromatic, erumpent to sometimes almost prominent ascomata with different size ranges. Due to the lack of sufficient molecular data spanning a wide range of morphodemes, we have not attempted to split this taxon further, after recognizing separate lineages forming flat, linear pseudostromata, such as A. flavostromatum (Aptroot & Cáceres 2016) and A. kunzei (see below), with an inspersed hamathecium, such as A. inspersaeneum (Lima et al. 2013; see below) and A. neoinspersum (Aptroot et al. 2016 a ), or with smaller or larger ascospores, such as A. flavum (Aptroot & Cáceres 2016) and A. megaeneum (Flakus et al. 2016).

New country record. Ecuador: Zamora-Chinchipe: Cordillera Numbala, Reserva Biologica San Francisco, 2004, Sipman & Mandi 53115 (B).

Astrothelium alboverrucum (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816653

Laurera alboverruca Makhija & Patw., Mycotaxon 31: 571 (1988); type: India, Andaman Islands, South Andaman, Port Mout, 14 ii 1985, Patwardhan, Nagarkar & Sethy AMH 85.36 (ABL!—isotype).

(Fig. 35G)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·9–1·3 mm diam., prominent, hemispherical, basally to laterally covered by thallus but upper part whitish, rough, with narrow, blackish ostiolar spot. Hamathecium clear, yellow, IKI−. Ascospores 6–8 per ascus, densely muriform, fusiform, (85–)100–170×23–33 µm, with distinctly thickened median septum, yellow, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (reported from India).

Astrothelium amazonum (R. C. Harris) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816654

Cryptothelium amazonum R. C. Harris, Acta Amazon. (Suppl.) 14: 65 (‘1984’) [1986]; type: Brazil, Amazonas, Reserva Biológica de Campina, Dumont 107 (NY!—isotype).

Verrucaria aurantia Eschw. in Martius, Icon. Sel. Pl. Crypt. 2: 15 (1828) nom. illeg., non (Pers.) Wibel (1799); type: Brazil, Martius s. n. (M—holotype, not seen).

(Fig. 37I)

Thallus corticate, light olive-green to yellowish, uneven to somewhat bullate and often pseudogall-inducing.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, not pseudostromatic, 0·8–1·2 mm diam., erumpent to prominent, wart-shaped to conical, covered by thallus except for papilliform, reddish to brown-black ostiole, internally with yellow-orange, K+ purple, UV+ red pigment. Hamathecium clear. Ascospores 8 per ascus, muriform, oblong-fusiform, 75–95×23–28 µm, median septum thickened, hyaline, IKI−.

Chemistry. Thallus UV−, K−, ascomata internally with yellow-orange, K+ purple, UV+ red anthraquinone.

Distribution. Neotropical (previously reported from Guyana and Brazil).

New country record. Venezuela: Amazonas: Alto Orinoco, 15 km W of Esmeralda, 110 m, 1997, Hafellner & Komposch 178-3-16 (GZU).

Astrothelium ambiguum (Malme) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816655

Laurera ambigua Malme, Ark. Bot. 19(1): 24 (1924).—Meristosporum ambiguum Malme (1924) nom. inval.; type: Brazil, Matto Grosso, Cuyabá, Malme 2009 (S!—lectotype, designated here; BM!—isolectotype).

(Fig. 33F)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·4–0·5 mm diam., erumpent to prominent, hemispherical, mostly exposed, brown-black. Hamathecium inspersed, IKI−. Ascospores 8 per ascus, muriform, fusiform, 45–60×14–20 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Astrothelium andamanicum (Makhija & Patw.) Aptroot comb. nov.

MycoBank No.: MB 816656

Cryptothelium andamanicum Makhija & Patw., J. Econ. Tax. Bot. 10: 498 (‘1987’) [1988].—Laurera bispora D. D. Awasthi, Lichenology in Indian Subcontinent: 15 (2000) non Laurera andamanica D. D. Awasthi (1991); type: India, Andaman Islands, North Andaman, Tugapur Range, Pathat Tikri, 19 xii 1985, Nagarkar & Sethy AMH 85.2323 (ABL!—isotype).

(Fig. 38J)

Thallus corticate, olive-green, smooth to uneven.

Ascomata pleurothelioid, with eccentric, separate ostioles, solitary, 0·6–1·0 mm diam., erumpent to prominent, partially covered by thallus but becoming exposed and black. Hamathecium clear. Ascospores 2 per ascus, densely muriform, fusiform, 80–130×23–36 µm, hyaline, IKI−, without distinctly thickened median septum.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (reported from India).

Astrothelium annulare (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816657

Pyrenula annularis Fée, Essai Crypt. Écorc.: 73 (1824).—Verrucaria annularis (Fée) Spreng., Syst. Veget. 4(1): 245 (1827).—Trypethelium annulare (Spreng.) Mont., Ann. Sci. Nat., Bot., sér. 2, 19: 71 (1843).—Pseudopyrenula annularis (Spreng.) Müll. Arg., Flora 68: 331 (1885); type: South America, “corticis annosas Cinchonea lancifoliae” (M!—isotype).

Verrucaria exasperata Zenker, in Goebel & Kunze, Pharmazeut. Waarenkunde 1(3): 183 (1827).—Trypethelium exasperatum (Zenker) Zahlbr., Catal. Lich. Univ. 1: 492 (1922); type: South America, s. col. (lost).

Trypethelium annulare var. detrusum Mont., Ann. Sci. Nat., Bot. sér. 2, 19: 71 (1843); type: French Guiana, Leprieur s. n. (PC-Mont—syntypes, not seen).

Verrucaria myriomma Nyl., Ann. Sci. Nat. Bot. sér. 5, 7: 348 (1867).—Pseudopyrenula myriomma (Nyl.) Müll. Arg., Flora 66: 248 (1883); type: Colombia, Pie de Cuesta, Lindig 98 (H-Nyl 1117!—holotype; BR—isotype).

(Fig. 21A)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to confluent, not pseudostromatic, 0·6–0·8 mm diam., immersed-erumpent, covered by thallus except for the dark ostiole surrounded by a narrow, whitish rim. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 62–80×20–25 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (reported from Costa Rica, Colombia, French Guiana).

Discussion. According to Hekking & Sipman (1988), Verrucaria exasperata Zenker is conspecific with Trypethelium annulare var. detrusum Mont. and hence both are included here in the synonymy of Astrothelium annulare.

Astrothelium aurantiacum (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816658

Laurera aurantiaca Makhija & Patw., Mycotaxon 31: 572 (1988); type: India, Karnataka, Agumbe-Shringera road, Patwardhan & Nagarkar s.n. (AMH—holotype, not seen).

(Fig. 33K)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·8–1·3 mm diam., strongly prominent to sessile, hemispherical to barrel-shaped with flattened top, covered by thallus but ostiolar area greyish. Hamathecium inspersed with red, K+ purple droplets. Ascospores 8 per ascus, densely muriform, ellipsoid, 115–135×26–33 µm, with distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected except for red anthraquinone in hamathecium.

Distribution. Eastern palaeotropical (previously reported from India and Papua New Guinea).

New country record. Malaysia: Sarawak: Gunong Mulu National Park, 1978, Coppins 5139 (ABL, E).

Astrothelium auratum (R. C. Harris) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816659

Laurera aurata R. C. Harris, Acta Amazon. (Suppl.) 14: 67 (‘1984’) [1986]; type: Brazil, Acre, km 12 on road to Porto Velho, Lowy 961 (NY!—isotype).

(Fig. 32D)

Thallus corticate, olive-green to brownish, uneven to verrucose.

Ascomata trypethelioid, with apical ostioles, distinctly pseudostromatic; pseudostromata 1–2 mm diam., prominent to sessile, each with 5–15 ascomata, with yellow-orange pigment. Hamathecium inspersed, IKI−. Ascospores 8 per ascus, muriform, ellipsoid, 60–75×20–30 µm, without distinctly thickened median septum, hyaline, IKI+ violet.

Chemistry. Thallus UV−, K−; pseudostromata with yellow-orange, K+ purple, UV+ red anthraquinone.

Distribution. Neotropical; reported from Brazil.

New country records. Venezuela: Amazonas: Alto Orinoco, 15 km W of Esmeralda, 110 m, 1997, Hafellner & Komposch 178-5-43 (GZU).—Guyana: Potaro-Siparuni: surroundings of Paramakatoi Village, 800 m, 1996, Sipman 41197 p.p. (B).

Astrothelium aureomaculatum (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816660

Pseudopyrenula aureomaculata Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 207 (1890).—Trypethelium aureomaculatum (Vain.) Zahlbr., Catal. Lich. Univ. 1: 488 (1922); type: Brazil, Minas Gerais, Caraça, Vainio s.n. (TUR-Vain 30749!—holotype; BM!—isotype; Vainio, Lich. Bras. 1473).

(Fig. 23D)

Thallus ecorticate, light brownish to greenish grey, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 0·5–1·0 mm broad, immersed to erumpent, with several joined ascomata, forming irregular groups or lines, with yellow pigment cover except for blackish ostiolar areas. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 28–35×10–15 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone; pseudostromata with yellow, K+ purple, UV+ red anthraquinone.

Discussion. This species is removed here from synonymy with Astrothelium versicolor, from which it differs substantially in thallus and ascoma morphology, the latter having a thick, bullate, whitish thallus with the joined ascomata dispersed.

Distribution. Neotropical (reported from Brazil).

Astrothelium basilicum (Kremp.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816661

Verrucaria basilica Kremp., Bol. Acad. Nac. Ci. Republ. Argentina 3: 126 (1879).—Pseudopyrenula basilica (Kremp.) Müll. Arg., Flora 72: 68 (1889).—Porina basilica (Kremp.) Zahlbr., Catal. Lich. Univ. 1: 367 (1922).—Trypethelium basilicum (Kremp.) R. C. Harris, Lichenogr. Thomsoniana: 141 (1998); type: Uruguay, Lorentz & Hieronymus s. n. (M!—lectotype; Harris, Lichenogr. Thomsoniana: 141, 1998; BM!—isolectotype).

(Fig. 21K)

Thallus corticate, olive-green, verrucose-bullate.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·8–1·5 mm diam., erumpent to prominent, covered by thallus except for blackish ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 11–19-septate, fusiform, 140–190×20–25 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (reported from Uruguay).

Astrothelium bicolor (Taylor) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816662

Trypethelium bicolor Taylor, in Hooker, London J. Bot. 6: 157 (1847).—Bathelium bicolor (Taylor) C.W. Dodge, Nova Hedwigia, Beih. 12: 20 (1964); type: Africa (BM!—isotype).

?Trypethelium scoria var. convexum Nyl., Mém. Soc. Acad. Maine-et-Loire 4: 74 (1858).—Trypethelium mastoideum var. convexum (Nyl.) Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 30(3): 12 (1888): type: “Crotone cascarillae” (PC—not seen).

Trypethelium polychroum Müll. Arg., Bot. Jahrb. Syst. 6: 391 (1885); type: Cuba, Wright s. n. (G!—lectotype, designated here; BM!—isolectotype; Müller, Verr. Cub. 178).

?Trypethelium scoria var. janeirense Zahlbr., Sitzungsber. K. Akad. Wiss., math.-naturw. Classe 111(1): 369 (1902); type: Brazil, Rio de Janeiro, von Höhnel 145 (W—holotype, not seen).

?Trypethelium formosanum Zahlbr., Feddes Repert. 31: 204 (1933); type: Taiwan, Taihoku, Soozan, Asahina 311 (W—holotype, not seen).

?Trypethelium boninense Kurok., Bull. Nat. Sci. Mus. Tokyo 12: 691 (1969); type: Japan, Ogasawara-shoto (Bonin Islands), Inoue 18960 (TNS—not seen).

(Fig. 18J)

Thallus corticate, green to yellowish or brown, smooth to somewhat bullate.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·7–1·5 mm broad, erumpent to prominent, rounded to irregular but not becoming confluent or reticulate, covered by thallus but often slightly paler than the surrounding thallus or sometimes whitish. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 15–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical (previously reported from, e.g. Mexico, Guadeloupe, Puerto Rico, Cuba, Dominican Republic, Netherlands Antilles, Costa Rica, Nicaragua, El Salvador, Ecuador, Bolivia, Guyana, Brazil, Argentina, Seychelles, India, China, Hong Kong, Taiwan, Japan, Malaysia, Philippines, Indonesia, Papua New Guinea, Solomon Islands, Australia, New Caledonia, Fiji and Hawai’i).

Discussion. This reinstated taxon belongs in the Astrothelium nitidiusculum complex, in which the epithet bicolor is the oldest for specimens with a clear hamathecium and ascomata arranged in pseudostromata; previously, this and other names were included in a wide species concept of Trypethelium nitidiusculum. Astrothelium bicolor appears to be the most common species in the complex. Due to the fact that the types of some presumed synonyms could not be tested for hymenium inspersion, some of these may eventually turn out to be conspecific with other species in this complex, such as A. nitidiusculum s. str. or the inspersed A. scoria.

New or confirmed country records. Mexico: Chiapas: Ocozocoautla, 1994, Wolf & Sipman 2205 (B).—Guadeloupe: Marie Galante, 1992, Vivant (ABL).—Puerto Rico: Mayagüez: Reserva Forestal Maricao, 1989, Sipman 25757 (B).—Dominican Republic: Distrito Nacional: Sierra Prieta, 1987, Harris 20151 (ABL, NY).—Netherlands Antilles: St. Eustatius: Quill, 2008, Sipman 56749 (B). Saba: The Bottom, 2007, Sipman 54960 (B).—Costa Rica: Puntarenas: Parque Nacional Carara, 2002, Sipman 48382l (ABL, B). Nicaragua: Rio San Juan: El Castillo, 2001, Breuss 18994 (LI).—El Salvador: Ahuachapán: Parque Nacional El Imposible, 1998, Sipman et al. 44882 (B).—Ecuador: Esmeraldas: San Lorenzo, 1982, Aptroot & Hensen 11022 (ABL). Pichincha: 19 km S of Santo Domingo, 1982, Aptroot & Hensen 11251 (ABL).—Guyana: Upper Takutu: Kusad Mountain, 1992, Sipman 57532 (B).—Brazil: Minas Gerais: Caraça, 1997, Sipman 40779 (B).—Argentina: Misiones: Iguazú, Ferraro et al. 10669 (ABL, CTES).—Seychelles: Mahé: Old Mission, 1973, Norkett 16442 (ABL, B).—India: West Bengal: Sundarbans, Chamta, 2003, Jagadeesh Ram 13547 (ABL).—China: Yunnan: Xishuangbanna, Menglun, 2002, Aptroot 57141 (ABL).—Hong Kong: New Territories: Tai Po Kau, 2000, Aptroot 48748 (ABL).—Malaysia: Pahang: Fraser’s Hill, 1964, Degelius s. n. (UPS).—Philippines: Luzon: Sorsogon, Irosin, 1915, Elmer 14659 (B).—Indonesia: Java: Bogor, Gunung Salak, 1991, Sipman & Tan 30065 (B).—Papua New Guinea: Madang: 10 km W of Brahman Mission, 1995, Sipman 38792 (B).—Solomon Islands: Guadalcanal Island: 1965, Hill 9215 (BM).—Australia: Queensland: 8 km W of Ravenshoe, 1984, Streimann 30176 (B).—New Caledonia: Ouenarou, 1970, Degelius P127 (UPS).—Fiji: Viti Levu: Nadarivatu, 1968, Degener & Degener 31814o (B).—Hawai’i: Hawai’i: Pahoa, 1969, Degener & Degener 32285b (B).

Astrothelium buckii (R. C. Harris) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816663

Trypethelium buckii R. C. Harris, Acta Amazon. (Suppl.) 14: 72 (‘1984’) [1986]; type: Brazil, Amazonas, along Rio Negro, 100 km NW of Manaus, Paraná Concelção, Buck 2157 (NY!—isotype).

(Fig. 14J)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent but not pseudostromatic, 0·6–0·9 mm diam., erumpent, covered with thallus but upper part reddish brown and ostiolar area blackish, internally with hyaline or red, K+ red crystals. Hamathecium inspersed. Ascospores 8 per ascus, 3-septate, fusiform, (32–)37–47×14–16 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−; ascomata internally K+ red, with anthraquinone.

Distribution. Neotropical (reported from Costa Rica, Venezuela, Brazil).

Discussion. This species is taken out of the synonymy with Astrothelium cartilagineum as suggested by Aptroot et al. (2008), differing from the latter by the inspersed hamathecium.

Astrothelium calosporum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816664

Pseudopyrenula calospora Müll. Arg., Bot. Jahrb. Syst. 6: 409 (1885).—Trypethelium calosporum (Müll. Arg.) R. C. Harris, Lichenogr. Thomsoniana: 141 (1998); type: Cuba, Wright s. n. (G!—holotype; BM!—isotype; Müller, Verr. Cub. 234).

Pseudopyrenula calospora var. rhodocheila Vain., Proc. Amer. Acad. Arts Sci. 58: 145 (1923); type: Trinidad, La Selva Valley, Thaxter 63 (S!—isotype).

(Fig. 17I)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·7–0·9 mm diam., immersed-erumpent, completely covered by thallus except for ostiole furnished with pink pigment. Hamathecium clear. Ascospores 8 per ascus, 14–18-septate, fusiform, 110–150×17–22 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−; ostiole K+ red, with anthraquinone.

Distribution. Neotropical (reported from Cuba and Trinidad).

Astrothelium campylocartilagineum Aptroot & Lücking nom. nov.

MycoBank No.: MB 816665

Campylothelium cartilagineum Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 195 (1890) non Astrothelium cartilagineum (Fée) Aptroot (see below); type: Brazil, Minas Gerais, Sitio, Vainio s. n. (TUR-Vain 30835!—holotype; BM!—isotype; Vainio, Lich. Bras. Exs. 1145).

(Fig. 39C)

Thallus corticate, olive-green, verrucose-bullate.

Ascomata pleurothelioid, with eccentric, separate ostioles, solitary, 0·7–1·0 mm diam., erumpent to prominent, covered by thallus but upper part often irregularly exposed and blackish. Hamathecium clear. Ascospores 2–4 per ascus, densely muriform, fusiform, 120–175×38–50 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (reported from Costa Rica and Brazil).

Discussion. This species is not synonymous with Heufleria chlorogastrica, as suggested by Harris (1986), since the type of the latter appears to belong in the genus Phyllobathelium.

Astrothelium cartilagineum (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816668

Pyrenula cartilaginea Fée, Essai Crypt. Écorc.: 76 (1824).—Trypethelium cartilagineum (Fée) Aptroot, in Aptroot et al., Biblioth. Lichenol. 98: 134 (2008) non Campylothelium cartilagineum Vain. (1890; see above); type: South America, “in America ad corticem Cinchonae lancifoliae et Portlandiae hexonae (quina nova)” (L!—isotype).

Thallus corticate, brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·4–0·5 mm diam., immersed to erumpent, covered by thallus except for dark ostiole surrounded by pale rim, internally with hyaline or red, K+ red crystals. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, (32–)37–47×14–16 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−; ascomata internally K+ red, with anthraquinone.

Distribution. Neotropical; previously reported from Trinidad, Costa Rica, French Guiana, Venezuela, and Brazil.

Discussion. This species is most similar to Astrothelium buckii, but differs by the clear hymenium.

New country record. Guyana: Upper Takutu: Kuyuwini Landing, 1992, Sipman 57073 (B); Karanambo ranch, 1992, Sipman 57280 (B).

Astrothelium cecidiogenum (Aptroot & Lücking) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816669

Cryptothelium cecidiogenum Aptroot & Lücking in Aptroot et al., Biblioth. Lichenol. 98: 57 (2008); type: Costa Rica, Alajuela, Volcán Tenorio National Park, Pilón Biological Station, Sipman 51911 (B!—holotype; INB-4055132!—isotype).

(Fig. 39K & L)

Thallus corticate, olive-green, smooth to uneven, pseudogall-inducing.

Ascomata mostly astrothelioid, several chambers joined with eccentric, fused ostioles, rarely pleurothelioid and with separate ostioles; joined ascomata diffusely pseudostromatic; pseudostromata 1–2 mm broad, prominent, confluent and forming reticulate or brain-like lines, covered with thallus except for the dark ostioles with irregular, whitish rim; pseudostromata 10–30 mm long and 5–10 mm wide, individual ascomata 0·7–1·2 mm diam. Hamathecium clear. Ascospores 8 per ascus, densely muriform, ellipsoid, 125–175×30–40 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution: Neotropical (previously reported only from Costa Rica).

New country record. Brazil: Rio Grande do Sul: Maquiné, 2006, Koch s. n. (ABL, HAS).

Astrothelium ceratinum (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816670

Pyrenula ceratina Fée, Essai Crypt. Écorc. Suppl.: 77 (1837).—Verrucaria ceratina (Fée) Nyl., Acta Soc. Sci. Fenn. 7(2): 491 (1863).—Pseudopyrenula ceratina (Fée) Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 30: 29 (1888).—Trypethelium ceratinum (Fée) R. C. Harris, Lichenogr. Thomsoniana: 141 (1998); type: Peru, “ad corticem cinchonarum” (G!—lectotype; Harris, Lichenogr. Thomsoniana: 141 (1998)).

(Fig. 19E)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·3–0·4 mm diam., erumpent, covered by thallus but upper portion darker or greyish. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 21–28×9–12 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Peru).

New country records. Guyana: Upper Mazaruni: N-slope of Mount Roraima, 1985, Sipman & Aptroot 18767 (Lichenotheca Latinoamericana 100, ABL).—Ecuador: Zamora-Chinchipe: Estacion San Francisco, 2004, Sipman 52735 (B).

Astrothelium chapadense (Malme) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816671

Laurera chapadensis Malme, Ark. Bot. 19(1): 23 (1924).—Meristosporum chapadense Malme (1924), nom. inval; type: Brazil, Matto Grosso, Santa Anna da Chapada, Malme 2513 (S!—holotype; S!—isotype).

(Fig. 36A)

Thallus corticate, olive-green to brownish yellow, uneven to verrucose or almost squamulose.

Ascomata trypethelioid, with apical ostioles, irregularly confluent to pseudostromatic; pseudostromata 0·8–2·0 mm broad, prominent to sessile, dark brown, forming irregular groups or lines. Hamathecium clear. Ascospores 8 per ascus, densely muriform, ellipsoid, (60–)70–100(–130)×20–30 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (reported from Costa Rica and Brazil).

Discussion. Astrothelium chapadense strongly resembles a species of Bathelium, but is included here in Astrothelium due to the lack of an internal anthraquinone pigment.

Astrothelium chrysoglyphum (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816672

Thelenella chrysoglypha Vain., Hedwigia 38: 258 (1899).—Laurera chrysoglypha (Vain.) Zahlbr., Catal. Lich. Univ. 1: 503 (1922); type: Guadeloupe, Parnasso, Duss 564 (TUR-Vain 31044!—holotype).

(Fig. 32L)

Thallus corticate, olive to yellowish but covered by orange pigment, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, aggregate to diffusely pseudostromatic, pseudostromata 1–2 mm broad, erumpent, covered by thallus and orange pigment except for blackish ostiolar area. Hamathecium clear. Ascospores 2 per ascus, densely muriform, fusiform, 80–150×25–40 µm, without distinctly thickened median septum, hyaline, IKI− or IKI+ brownish.

Chemistry. Thallus and ascomata UV+ red, K+ purple, with anthraquinone.

Distribution. Neotropical (reported from Guadeloupe).

Astrothelium chrysostomum (Kremp.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816673

Trypethelium chrysostomum Kremp., Nouvo Giorn. Bot. Ital. 7: 57 (1875); type: Malaysia, Sarawak, Beccari 50 (M!—lectotype, designated here following annotation by R. C. Harris).

Trypethelium leucostomum Kremp., Nouvo Giorn. Bot. Ital. 7: 57 (1875) non (Nyl.) C. W. Dodge (1953); type: Malaysia, Sarawak, Beccari 188 (M!—lectotype, designated here; M!—isolectotype).

(Fig. 23H & I)

Thallus corticate, olive-brown, uneven-rugose.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata solitary to confluent, 0·7–1·0 mm diam., immersed in thick thallus folds, covered except pale ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 3(–5)-septate, fusiform, 32–52×10–17 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−. TLC: lichexanthone.

Distribution. Eastern palaeotropical (Malaysia: Sarawak).

Discussion. This taxon was first believed to be conspecific with Astrothelium leucothelium but differs substantially in thallus morphology and also in its distribution.

Astrothelium cinereorosellum (Kremp.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816674

Trypethelium cinereorosellum Kremp., Nuovo Giorn. Bot. Ital. 7: 35 (1875); type: Sarawak, Beccari 48 (M!—lectotype; Makhija & Patw., J. Hattori Bot. Lab. 73: 194, 1993).

Verrucaria antoniae Kremp., Nuovo Giorn. Bot. Ital. 7: 51 (1875).—Pyrenula antoniae (Kremp.) van Overeem-de Haas, Bull. Jard. Bot. Buitenzorg, Ser. 3, 4: 113 (1922).—Pseudopyrenula antoniae (Kremp.) Zahlbr., Catal. Lich. Univ. 1: 354 (1922); type: Sarawak, Beccari (M!—lectotype; Harris, Lichenogr. Thomsoniana: 140, 1998).

Trypethelium microstomum Makhija & Patw., J. Hattori Bot. Lab. 73: 201 (1993); type: India, Andaman Islands, South Andaman, Baratang, Bishmu Nala, 22 ii 1985, Patwardhan & Nagarkar AMH 85.663 (ABL!—isotype).

Trypethelium flavocinereum Makhija & Patw., J. Hattori Bot. Lab. 73: 198 (1993); type: India, Kerala, Cardamom Hills, Munnar-Devicolam road, 25 January 1976, Patwardhan & Prabhu AMH 76.672 (ABL!—isotype).

(Fig. 12D & E)

Thallus corticate, light greenish grey to yellowish or whitish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary or forming irregular groups or lines but not distinctly pseudostromatic, 0·3–0·5 mm diam., erumpent to prominent, covered by thallus except dark ostiole. Hamathecium inspersed. Ascospores 8 per ascus, (5–)7–9-septate, fusiform, (33–)40–50(–73)×8–14 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone).

Distribution. Palaeotropical; previously reported from India, Thailand, Borneo, Papua New Guinea, and Australia.

New country record. Indonesia: Java: Cibodas, 1450 m, 1939, Groenhart 1942 (ABL, L).

Astrothelium cinereum (Müll. Arg.) Aptroot & Lücking comb. et stat. nov.

MycoBank No.: MB 816675

Heufleria praetervisa var. cinerea Müll. Arg., Flora 68: 251 (1885).—Cryptothelium praetervisum var. cinereum (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 522 (1922); type: French Guiana, Leprieur 41 (G!—lectotype, designated here).

(Fig. 39G)

Thallus corticate, olive-brown, uneven to coarsely bullate-folded.

Ascomata mostly astrothelioid, several chambers joined with eccentric, fused ostioles, rarely pleurothelioid and with separate ostioles; joined ascomata pseudostromatic, pseudostromata 0·7–1·5 mm diam., erumpent to prominent, forming irregular lines, covered by thallus except for dark ostiole surrounded by whitish rim. Hamathecium clear. Ascospores 8 per ascus, densely muriform, fusiform, 33–52×16–22 µm, without thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from French Guiana).

Notes. This taxon is very similar to Astrothelium praetervisum but appears to lack the thin orange pigment characteristic of the latter.

New country record. Guyana: Upper Mazaruni: Paruima Mission, 1997, Sipman 39819 (B). Upper Takutu: Kuyuwini Landing, 1992, Sipman 57079 (B).

Astrothelium cinnamomeum (Eschw.) Müll. Arg.

Flora 68: 270 (1884).—Pyrenastrum cinnamomeum Eschw., in Martius, Icon. Sel. Pl. Crypt. 2: 18, t. 9, fig. 1 (1828); type: Brazil, Bahia, Caetité, Martius s. n. (M!—holotype).

Astrothelium conicum Eschw., in Martius, Syst. Lich.: 18 (1833); Eschweiler, Syst. Lich.: 26 (1824), nom. nud.—Heufleria conica (Eschw.) Trevis., Spighe e Paglie: 19 (1853); type: Brazil, Martius s. n. (M!—holotype).

Astrothelium minus Müll. Arg., Bot. Jahrb. Syst. 6: 382 (1885); type: Cuba, Wright s. n. (G!—holotype; NY!—isotype; Müller, Verr. Cub. 235).

(Fig. 26I–L)

Thallus corticate, light olive-green to yellowish, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic, pseudostromata 0·4–0·7 mm diam., erumpent to prominent, conical with flattened top, covered by thallus and upper portion with yellow-orange, K+ purple, UV+ red pigment, area beneath the upper portion often whitish. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 23–30×6–10 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata with yellow to orange, K+ purple, UV+ red anthraquinone.

Distribution. Pantropical (previously reported from the USA, Mexico, Cuba, Trinidad, Tobago, Costa Rica, Guyana, French Guiana, Venezuela, Brazil, Seychelles, Hong Kong, Borneo and Australia).

Discussion. The synonymy of Astrothelium conicum (the type species of the genus) with A. cinnamomeum is confirmed by combining the observations of Müller (1884: 270), who observed the ascospores, and Harris (1986: 59), who observed no ascospores but saw the negative reaction with UV.

New country record. China: Yunnan: Xishuangbanna, Menglun, 2002, Aptroot 57255, 57026, 57267 (all ABL).

Astrothelium coccineum Córdova-Chávez et al.

In Córdova-Chávez et al., Cryptog., Mycol. 35: 159 (2014); type: Mexico, Veracruz, La Cortadure, Coatepec, 2062 m, 2013, Córdova-Chávez 464 (XAL!—holotype; ABL!—isotype).

(Fig. 14K)

Thallus corticate, olive-brown, verrucose-bullate.

Ascomata trypethelioid, with apical ostioles, distinctly pseudostromatic; pseudostromata 0·7–1·5 mm broad, erumpent to prominent, rounded to irregular, covered by dark red pigment except for the blackish ostioles, internally with yellow pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, oblong-ellipsoid, 25–30×10–13 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV−, outside K+ purple, inside K+ crimson red. TLC: a red anthraquinone at Rf 7 and a yellow anthraquinone at Rf 2.

Distribution. Neotropical (reported from Mexico).

Astrothelium confluens (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816676

Heufleria confluens Müll. Arg., Flora 66: 243 (1883).—Cryptothelium confluens (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 521 (1922); type: Brazil, Rio de Janeiro, Apiahy, Puiggari 143 (G!—lectotype, designated here).

(Fig. 37F)

Thallus corticate, greyish green to pale yellowish, uneven to bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles, rarely pleurotremoid with separate ostioles; joined ascomata 1–3 mm diam., prominent, completely covered by thallus. Hamathecium clear. Ascospores 8 per ascus, densely muriform, fusiform, c. 130×20 µm, with distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata UV+ yellow, K−. TLC: lichexanthone on the ascomata.

Distribution. Neotropical (previously reported only from Brazil).

New country record. Guyana: Potaro-Siparuni: Kaieteur Falls National Park, 1996, Sipman 40482 (B).

Astrothelium consimile (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816677

Heufleria consimilis Müll. Arg., Bot. Jahrb. Syst. 6: 385 (1885).—Cryptothelium consimile (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 521 (1922); type: Cuba, Wright s. n. (G!—holotype).

(Fig. 36L)

Thallus corticate, olive-green yellowish, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles, rarely pleurotremoid with separate ostioles; joined ascomata dispersed to confluent or diffusely pseudostromatic, 0·8–1·2 mm diam., immersed-erumpent, covered by thallus but upper portion whitish with blackish ostiole area. Hamathecium clear. Ascospores 2 per ascus, densely muriform, fusiform, 85–130×23–36 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. No reactions or substances in the type material; however, the specimen from Brazil has the medulla UV+ yellow.

Distribution. Neotropical (previously reported from Cuba).

New country record. Brazil: Rondônia: Porto Velho, Parque Natural Municipal, 2012, Cáceres & Aptroot 15568 (ABL, ISE).

Astrothelium crassum (Fée) Aptroot

In Aptroot et al., Biblioth. Lichenol. 98: 44 (2008).—Trypethelium crassum Fée, Essai Crypt. Écorc.: 66 (1824).—Pyrenodium crassum (Fée) Fée, Essai Crypt. Écorc. Suppl.: 69 (1837); type: South America, “ad ramae Crotonis cascarillae” (G! —lectotype; L!—isolectotype).

Trypethelium clandestinum Fée, Essai Crypt. Écorc.: 68 (1824).—Polyblastia clandestina (Fée) Trevis., Spighe e Pagie: 15 (1853), non Jatta (1900).—Astrothelium clandestinum (Fée) Nyl., Mém. Soc. Sci. Nat. Cherbourg 5: 141(1857).—Pyrenastrum clandestinum (Fée) Müll. Arg., Bot. Jahrb. Syst. 6: 386 (1885); type: South America, “Cinch. jaune” (G—holotype, not seen; L!—isotype).

Astrothelium confusum Müll. Arg., Flora 68: 247 (1885); type: Colombia, Rio Magdalena, Lindig 141 (G!—holotype; BM!, BR—isotypes).

(Fig. 29A & B)

Thallus corticate, olive-green to yellowish brown, rather thick, uneven to bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to confluent, 0·3–0·4 mm diam., erumpent, covered by thallus but upper portion whitish and ostiolar area blackish. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 21–27×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical; previously reported from Costa Rica, Guyana, Venezuela, Brazil, India, New Caledonia and Papua New Guinea.

New country records. Puerto Rico: Ponce: Caribbean National Forest, 1989, Aptroot & Aptroot 25523 (ABL).—Colombia: Amazonas: Araracuara, opposite Isla Morrocoy, 240 m, 1988, Sipman & Duivenvoorden 28502 (B).—Ecuador: Los Rios: 45 km S of Quevedo, 1982, Aptroot & Hensen 10422 (ABL).—Madagascar: Tamatave: Andasibe (Périnet), 1984, Aptroot & Hensen 13396 (ABL).—Indonesia: Java: South Semeru Lands, Tempur Sewa, 1939, Groenhart 1679, 2595 (ABL, L).

Astrothelium croceum Malme

Ark. Bot. 19(1): 12 (1924); type: Brazil, Matto Grosso, Serra da Chapada, Malme, 26 vii 1984 (S!—holotype).

(Fig. 27A–C)

Thallus corticate, light olive-green to yellowish, yellowish grey, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to confluent, 0·4–0·7 mm diam., erumpent, covered by thallus and yellow-orange, K+ purple, UV+ red pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 30–35×10–12 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−, without lichexanthone or pigment; ascomata with yellow-orange, K+ purple, UV+ red anthraquinone.

Distribution. Neotropical (reported from Brazil).

Astrothelium deforme (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816678

Trypethelium deforme Fée, Ann. Sci. Nat. 23: 454 (1831).—Meissneria varia Fée, Essai Crypt. Écorc. Suppl.: 66 (1837).—Bathelium deforme (Fée) Trevis., Spighe e Paglie: 20 (1853).—Trypethelium varium (Fée) Nyl., Mém. Soc. Acad. Maine-et-Loire 4: 78 (1858).—Bathelium varium (Fée) Trevis., Flora 44: 21 (1861).—Laurera varia (Nyl.) Zahlbr., in Engler & Prantl, Natürl. Pflanzenfamil. 1(1*): 71 (1903); type: Malaysia, Sarawak, Beccari (G-G00128943!—lectotype, designated here).

(Fig. 32H)

Thallus corticate, olive-green, uneven-verrucose.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·5–1·0 mm diam., erumpent, covered by thallus but upper portion blackish, internally with orange, K+ purple pigment. Hamathecium clear. Ascospores 8 per ascus, muriform, fusiform, 90–105×26–36 µm, without thickened median septum, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata UV−, externally K−, internally K+ purple.

Distribution. Eastern palaeotropical (reported from Malaysia: Sarawak).

Discussion. Fée later replaced his earlier epithet deforme by varia because he found the species to be variable rather than deformed; however, according to the retroactive nomenclatural rules, his earlier epithet has to be taken up.

Astrothelium defossum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816679

Heufleria defossa Müll. Arg., Flora 68: 250(1885).—Cryptothelium defossum (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 521 (1922) non Campylothelium defossum Müll. Arg. (1891); type: French Guiana, Leprieur 168 (G!—holotype).

(Fig. 37A–C)

Thallus corticate, olive-green to light greyish green, uneven to bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles, rarely pleurotremoid with separate ostioles; joined ascomata dispersed to confluent or diffusely pseudostromatic, 0·8–1·3 mm diam., erumpent, covered by thallus but upper, often papilliform portion whitish with blackish ostiole. Hamathecium clear. Ascospores 8 per ascus, densely muriform, fusiform, 100–150×30–45 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata UV+ yellow, K−. TLC: lichexanthone on the ascomata.

Distribution. Neotropical (previously reported from Venezuela and French Guiana).

New country record. Guyana: Upper Mazaruni: Paruima Mission, 1997, Sipman 39596 (B).

Astrothelium degenerans (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816680

Pseudopyrenula degenerans Vain., J. Bot. 34: 292 (1896).—Trypethelium degenerans (Vain.) Zahlbr., Catal. Lich. Univ. 1: 490 (1922).—Bathelium degenerans (Vain.) R. C. Harris, More Florida Lichens: 117 (1995); type: Dominica, Laudat, Elliott s. n. (TUR-Vainio 30782! —holotype; BM!—isotype).

(Fig. 16J–L)

Thallus corticate, olive-green to yellowish or brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, distinctly pseudostromatic; pseudostromata 1–2 mm diam., erumpent, forming irregular groups including numerous ascomata, dark brown, internally with yellow-orange, dusty granular pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 15–25×6–9 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV−, externally K−, internally K+ red, with anthraquinone.

Distribution. Pantropical (previously reported from Mexico, Dominica, Jamaica, Puerto Rico, Costa Rica, El Salvador, Venezuela, Galapagos and Thailand).

New country records. Netherlands Antilles: Saba: Maskehorn, 2007, Sipman 54934 (B).—Ecuador: Manabi: Puerto Lopez, 1982, Aptroot & Hensen 10573, 10594 (ABL).

Astrothelium diplocarpoides Müll. Arg.

Bot. Jahrb. Syst. 6: 384 (1885).—Astrothelium diplocarpoides Nyl. (1876) nom. nud; type: Cuba, Wright s. n. (H-Nyl 191!—holotype; BM!—isotype; Müller, Verr. Cub. 143).

(Fig. 25J)

Thallus corticate, light olive-green but with distinct, whitish to pale yellowish pruina (lichexanthone), smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 0·8–1·5 mm diam., prominent to almost sessile, covered by thallus and whitish pruina. Hamathecium inspersed. Ascospores 8 per ascus, 5–7-septate, fusiform, 80–85×20 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−. TLC: lichexanthone.

Distribution. Neotropical (reported from Cuba).

Astrothelium diplocarpum Nyl.

Flora 47: 618 (1864).—Heufleria diplocarpa (Nyl.) Müll. Arg., Bot. Jahrb. Syst. 6: 385 (1885).—Cryptothelium diplocarpum (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 521 (1922); type: Colombia, Pie de Cuesta, Lindig s. n. (H-Nyl 109!—holotype).

(Figs 26C & D, 37K)

Thallus corticate, olive-green to yellowish brown, uneven-bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 0·7–1·0 mm diam., immersed-erumpent, covered by thallus except for the papilliform, orange to brown, K+ red ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 9-septate to submuriform, fusiform, 90–110×22–28 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata with brown, K+ red anthraquinone.

Distribution. Neotropical (previously reported from the USA, Costa Rica, Colombia and Ecuador).

New country record. Venezuela: Bolivar: Cerro Guaiquinima, 1000 m, 1990, Sipman 26568 (B).

Astrothelium dissimilum (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816681

Trypethelium dissimilum Makhija & Patw., J. Hattori Bot. Lab. 73: 195 (1993); type: India, Meghalaya, Mawsyhram, 30 October 1977, Patwardhan & Nagarkar AMH 77.1169 (ABL!—isotype).

(Fig. 17C & D)

Thallus corticate, olive-green, uneven-verrucose.

Ascomata trypethelioid, with apical ostioles, confluent to pseudostromatic; pseudostromata 0·8–2·0 mm diam., prominent, dark brown, internally with yellow, K+ purple crystals. Hamathecium clear. Ascospores 8 per ascus, (3–)5(–9)-septate, fusiform, 25–40×6–9 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, externally K−; pseudostromata internally K+ red, with anthraquinone.

Distribution. Eastern palaeotropical (India and South Korea).

Discussion. The material from South Korea is tentatively assigned to this species, although the ascomata are smaller and less emergent (Fig. 17D).

Astrothelium effusum (Aptroot & Sipman) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816682

Laurera effusa Aptroot & Sipman in Aptroot et al., Biblioth. Lichenol. 98: 61 (2008); type: Costa Rica, Puntarenas, La Amistad International Park, Altamira Station, 1500–1600 m, Sipman 48034e (B!—holotype; INB-3944626!—isotype).

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·6–1·0 mm diam., immersed-erumpent, covered by thallus except dark ostiole surrounded by pale ring. Hamathecium inspersed. Ascospores 8 per ascus, muriform, ellipsoid, 80–100(–125)×28–48 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Costa Rica).

Astrothelium endochryseum (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816683

Pseudopyrenula endochrysea Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 206 (1890).—Trypethelium endochryseum (Vain.) Zahlbr., in Engler & Prantl, Natürl. Pflanzenfamil. 1(1*): 70 (1903).—Bathelium endochryseum (Vain.) R. C. Harris, More Florida Lichens: 117 (1995); type: Brazil, Minas Gerais, Caraça, Vainio s. n. (TUR-Vain 30793!—holotype; BM!—isotype; Vainio, Lich. Bras. 1157).

(Fig. 16B & C)

Thallus corticate, green to yellowish or brown, verrucose.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 1–2 mm diam., erumpent to prominent, dark brown, internally with yellow to orange, dusty granular pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 35–50×12–15 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV−, externally K−, internally K+ red, with anthraquinone.

Distribution. Neotropical (Brazil).

Astrothelium eustomum (Mont.) Müll. Arg.

Flora 68: 247 (1885).—Pyrenastrum eustomum Mont., Ann. Sci. Nat. Bot., sér. 2 19: 63 (1843); type: French Guiana, Leprieur 179 (PC!—holotype; G! —isotype).

Trypethelium celatum Stirt., Proc. Roy. Philos. Soc. Glasgow 13: 193 (1881); type: India, Assam, Lezhore, Watt (BM!—isotype).

Astrothelium acroleucum Malme, Ark. Bot. 19(1): 15 (1924); type: Brazil, Matto Grosso, Santa Anna da Chapada, Malme 2358 (S!—lectotype, designated here).

(Fig. 25D)

Thallus corticate, olive-green to greyish green, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to pseudostromatic, 0·4–0·8 mm diam., erumpent with flattened tops, with whitish to pale yellowish cover contrasting with the thallus. Hamathecium clear. Ascospores 8 per ascus, 3–5-septate, fusiform, 22–30×6–8 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone).

Distribution. Pantropical; previously reported from Costa Rica, Cuba, Guyana, French Guiana, Colombia, Venezuela, Brazil, Bolivia, India, Sri Lanka, Thailand and Australia.

Discussion. The three types cited above show the two extremes in terms of ascoma emergence and pseudostroma development, with Pyrenastrum eustomum and Trypethelium celatum showing the ascomata almost fully immersed in the thallus and mostly dispersed, and Astrothelium acroleucum showing erumpent pseudostromata.

New country records. Madagascar: Tamatave: Andasibe (Périnet), 1984, Aptroot & Hensen 13384 (ABL).—Papua New Guinea: Central: Varirata National Park, 1995, Sipman 38662 (B).—Solomon Islands: Guadalcanal Island: 1965, Hill 9119 (BM).

Astrothelium exostemmatis (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB816684

Bathelium exostemmatis Müll. Arg. Mém. Soc. Phys. Hist. Nat. Genève 30(3): 17 (1888); Laurera exostemmatis (Müll. Arg.) Zahlbr. in Engler & Prantl, Natürl. Pflanzenfamil. 1(1*): 71 (1903); type: Jamaica, “cort. Exostemmatis” (G!—holotype; M—isotype, not seen).

(Fig. 35L)

Thallus corticate, olive-brown to pale brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly clustered to diffusely pseudostromatic, 0·8–1·2 mm diam., when confluent pseudostromata up to 5 mm diam., immersed-erumpent, covered by thallus except dark ostiole surrounded by irregular whitish rim. Hamathecium clear. Ascospores 8 per ascus, muriform, ellipsoid, distinctly curved, 70–80×20–25 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances.

Distribution. Neotropical (previously reported from Jamaica).

New country record. Ecuador: Pastaza: Puyo, 1100 m, 1982, Aptroot & Hensen 10406 (ABL).

Astrothelium fallax Müll. Arg.

Bot. Jahrb. Syst. 6: 383 (1885).—Trypethelium pallescens Leight., Trans. Linn. Soc. London 27: 185 (1869) non Fée (1831); type: Sri Lanka, Thwaites 195 (BM!—lectotype, designated here; PC!, S!—isolectotypes).

(Fig. 29D & E)

Thallus corticate, olive-green to yellowish, uneven to strongly bullate, inducing pseudogall formation.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic, 0·8–1·5(–2·0) mm diam., prominent to sessile, covered by thallus except for the blackish ostiole area surrounded by a whitish ring, forming large warts with uneven to bumpy surface. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 30–39×10–13 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical; previously reported from Cuba, India and Sri Lanka.

New country record. Guyana: Upper Mazaruni: Mount Latipu, 1985, Sipman & Aptroot 19120 (ABL, B).

Astrothelium feei (C. F. W. Meissn.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816685

Trypethelium feei C. F. W. Meissn., in Fée, Ann. Sci. Nat., Bot. sér. 1 23: 442 (1831).—Trypethelium scoria var. feei (C. F. W. Meissn.) Trevis., Flora 44: 10 (1861).—Bathelium feei (C. F. W. Meissn.) Aptroot, in Aptroot et al., Biblioth. Lichenol. 98: 51 (2008); type: South America, “Cascarill.” (G!—holotype; L!—isotype).

Trypethelium mastoideum var. macerum Müll. Arg., Bot. Jahrb. Syst. 6: 390 (1885); type: Cuba, Müller, Verr. Cub. 153 (G!—lectotype, designated here; BM!—isolectotype).

(Fig. 16F–I)

Thallus corticate, olive-brown to olive-green or yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, distinctly pseudostromatic; pseudostromata 1–2 mm diam., erumpent, dark brown, internally with yellow-orange, dusty granular pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 18–25×5–9 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV−, externally K−, internally K+ yellow, with anthraquinone.

Distribution. Neotropical; previously reported from the USA, Costa Rica, Colombia, French Guiana and Galapagos.

Discussion. The difference in pseudostroma chemistry from the similar Astrothelium degenerans seems to warrant the distinction of the two species, but it is not correlated with distribution or any other character.

New country records. El Salvador: Ahuachapán: Parque Nacional El Imposible, 1999, Sipman & Bohnke 44912 (B).—Venezuela: Acosta, Aqua Salada, 1989, Kalb & Kalb s. n. (hb. Kalb).—Guyana: Marudi Mts, Aishalton, 1982, Stoffers et al. 251e (ABL).

Astrothelium ferrugineum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816686

Trypethelium ferrugineum Müll. Arg., Bot. Jahrb. Syst. 6: 392 (1885); type: Cuba, Wright s. n. (G!—lectotype, designated here; Müller, Lich. Cub. 593).

Trypethelium ferrugineum var. inornatum Müll. Arg., Bot. Jahrb. Syst. 6: 392 (1885); type: Cuba, Wright s. n. (G!—lectotype, designated here; Müller, Verr. Cub. 157).

(Fig. 26F & G)

Thallus corticate, olive-green but sparsely covered by orange pigment, uneven to shallowly verrucose.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 0·8–1·5 mm diam., prominent, covered by thallus and sparse orange pigment except for the blackish ostioles. Hamathecium clear. Ascospores 8 per ascus, 3-septate, 20–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV+ red, K+ purple, with anthraquinone.

Distribution. Neotropical (Cuba).

Distribution. This species has recently been synonymized with Trypethelium aeneum (Harris 1995), but it is kept separate here as no morphological intermediates occur and the world distribution and ecology seems to differ; this appears to be a very rare species.

Astrothelium floridanum Zahlbr. ex M. Choisy

In Choisy, Icon. Lich. Univ.: pl. 5 (1928).—Trypethelium floridanum (Zahlbr. ex M. Choisy) R. C. Harris, Acta Amazon. (Suppl.) 14: 73 (‘1984’) [1986]; type: USA, Florida, Rapp (W—holotype, not seen).

(Fig. 20A & B)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent but not distinctly pseudostromatic, 0·3–0·6 mm diam., erumpent, covered by thallus except for the blackish ostiolar area surrounded by a whitish rim. Hamathecium clear. Ascospores 8 per ascus, 3-septate, 35–45×14–16 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Pantropical, extending into temperate regions; previously reported from the USA, Costa Rica and Japan.

Discussion. The synonymization with Astrothelium marcidum by Aptroot et al. (2008) was based on a specimen in L that was apparently incorrectly thought to be an isotype of that taxon. The correct type has astrothelioid ascomata (see below).

New country records. Puerto Rico: Ponce: Caribbean National Forest, 1150 m, 1989, Aptroot & Aptroot 25608 (ABL).—Venezuela: Bolivar: Cerro Guaiquinima, 1000 m, 1990, Sipman 26683 (B).—Colombia: Amazonas: Araracuara, opposite Isla Mariñame, 240 m, 1988, Sipman & Duivenvoorden 28337 (B).

Astrothelium galligenum (Aptroot) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816687

Trypethelium galligenum Aptroot, Tropical Bryology 14: 29 (1998); type: Papua New Guinea, Central Province, Owen Stanley Range, Kagi Village, along Kokoda Trail towards Gap, Aptroot 39461 (B!—holotype; ABL!—isotype).

(Fig. 21B)

Thallus corticate, pale greenish, uneven to bullate, causing pseudogall formation.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·8–1·5 mm diam., erumpent, apically whitish and blackish ostioles surrounded by a brown rim. Hamathecium inspersed with yellow, K+ purple oil. Ascospores 8 per ascus, 7-septate, 50–60×12–17 µm, central septum euseptate and wider, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−; hamathecium internally K+ purple, with anthraquinone.

Distribution. Eastern palaeotropical (Papua New Guinea).

Astrothelium gigantosporum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816688

Bathelium gigantosporum Müll. Arg., Bot. Jahrb. Syst. 6: 394 (1885).—Laurera gigantospora (Müll. Arg.) Zahlbr., in Engler & Prantl, Natürl. Pflanzenfamil. 1(1*): 71 (1903); type: Cuba, Wright s. n. (G!—holotype).

(Fig. 35C & D)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic, pseudostromata 1–3(–5) mm diam., erumpent to prominent, covered by thallus except for dark ostioles surrounded by whitish rim. Hamathecium clear. Ascospores 2 per ascus, densely muriform, fusiform, 200–280×50–65 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Cuba and Costa Rica).

New country record. Ecuador: Zamora-Chinchipe: Cordillera Numbala, Reserva Biologica San Francisco, 2004, Sipman 52875, 52646 (B).

Astrothelium gigasporum R. C. Harris

Acta Amazon. (Suppl.) 14: 61 (‘1984’) [1986]; type: Brazil, Amazonas, along Rio Curicuriari, Cachoeira Piraiwara, Buck 2532 (ABL!, NY!—isotypes).

(Fig. 30K & L)

Thallus corticate, olive-green, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata diffusely pseudostromatic, pseudostromata 0·8–1·5 mm diam., prominent to sessile, covered by thallus but with strongly protruding, papilliform, brown ostioles. Hamathecium clear. Ascospores 8 per ascus, 19–22-septate, fusiform, 105–115×18–20 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical; previously reported from Guyana, Venezuela and Brazil.

New country records. Colombia: Amazonas: Araracuara, Villazul, 300 m, 1988, Sipman & Duivenvoorden 28498 (B).—French Guiana: Saül, 1988, Sipman 31687 (B).

Astrothelium grossoides Aptroot & Lücking nom. nov.

MycoBank No.: MB 816689

Trypethelium grossum Müll. Arg., Bot. Jahrb. Syst. 5: 139 (1884), non Astrothelium grossum Müll. Arg. (1888); type: Papua New Guinea, Naumann 409 (BM!—isotype).

(Fig. 12I)

Thallus corticate, olive-green to brownish, uneven-bullate, pseudogall-inducing.

Ascomata trypethelioid, with apical ostioles, dense to diffusely pseudostromatic, 0·4–0·5 mm diam., immersed, covered by thallus. Hamathecium inspersed. Ascospores 8 per ascus, 3-septate, 15–32×9–12 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−. TLC: lichexanthone.

Distribution. Eastern palaeotropical; reported from Papua New Guinea, where it is locally abundant, and New Caledonia.

Astrothelium grossum Müll. Arg.

Flora 71: 141 (1888); type: New Caledonia, Pancher, 1870 (G!—holotype).

(Fig. 23E)

Thallus corticate, greenish to yellowish, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to confluent but usually not pseudostromatic, 0·4–0·6 mm diam., erumpent to prominent, wart-shaped, covered by thallus and yellow-orange, K+ purple, UV+ red pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, 30–35×10–15 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone; ascomata with yellow to orange, K+ purple, UV+ red anthraquinone.

Distribution. Reported only from New Caledonia, from where we also observed additional specimens.

Discussion. This taxon corresponds to what has traditionally been named Astrothelium versicolor. The type of the latter has a very peculiar thallus morphology and hence we suspect that many collections identified with the name A. versicolor might represent A. grossum.

Astrothelium heterophorum Nyl.

Bull. Soc. Linn. Normand., sér. 2 2: 133 (1868); type: New Caledonia, Lifu, Thiébaut (H-NYL—holotype, not seen).

Trypethelium elmeri (Vain.) R. C. Harris, Lichenogr. Thomsoniana: 143 (1998).—Pseudopyrenula elmeri Vain., Ann. Acad. Sci. Fenn., ser. A 6: 354 (1921); type: Philippines, Luzon, Irosin, Sorsogon, Elmer 14655 (TUR-Vain 30812!—holotype; ABL!, G!—isotypes).

(Fig. 30E)

Thallus corticate, olive-green to yellowish, smooth to uneven.

Ascomata pleurothelioid and with separate ostioles or in part astrothelioid, several chambers joined with eccentric, fused ostioles; single or joined ascomata dispersed, 0·5–1·0 mm diam., immersed-erumpent, covered by thallus except for blackish ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 9–19-septate, fusiform, 60–80×12–17 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (Philippines and New Caledonia).

Astrothelium indicum (Upreti & Ajay Singh) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816690

Laurera indica Upreti & Ajay Singh, Bull. Jard. Bot. Nat. Belg. 57: 372 (1987) non Makhija & Patw. (1988); type: India, Bengal, Kurz 202 (H-Nyl 122!—holotype; BM!—isotype).

(Fig. 33G)

Thallus corticate, olive-green to brownish but whitish pruinose, uneven-rugose.

Ascomata trypethelioid with apical ostioles, more or less grouped but not distinctly pseudostromatic, groups of ascomata 0·8–1·2 mm diam., erumpent, covered by thallus and whitish pruina except for the blackish ostiolar area. Hamathecium inspersed. Ascospores (4–)8 per ascus, muriform, ellipsoid, (33–)50–70×13–17 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (India and Thailand).

Astrothelium infossum (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816691

Verrucaria infossa Nyl., Flora 69: 178 (1886).—Pseudopyrenula infossa (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 357 (1922).—Trypethelium infossum (Nyl.) R. C. Harris, Lichenogr. Thomsoniana: 144 (1998); type: São Tomé, Bom Successo, Moller s. n. (H-Nyl 1034!—holotype; BM!—isotype).

(Fig. 18G)

Thallus corticate, olive-brown to yellowish brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·3–0·4 mm diam., immersed, covered by thallus except for blackish ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 22–27×7–9 µm, hyaline, IKI+ violet.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Africa (São Tomé).

Discussion. A very inconspicuous species except for the amyloid ascospores, a rare feature in the family. Otherwise, the species falls within the Astrothelium nitidiusculum complex.

Astrothelium infuscatulum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816692

Trypethelium infuscatulum Müll. Arg., Bot. Jahrb. Syst. 6: 389 (1885).—Pseudopyrenula infuscatula (Müll. Arg.) Vain., Ann. Acad. Sci. Fenn., ser. A 6(7): 197 (1915); type: Cuba, Wright s. n. (G!—holotype; L!—isotype; Müller, Verr. Cub. 175).

(Fig. 18H & I)

Thallus corticate, olive-green to yellowish brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, distinctly pseudostromatic; pseudostromata 1–2 mm broad, erumpent, irregular to linear, with flattened surface, blackish brown except for a pale brown rim around the ostioles. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 25–30×7–10 µm, hyaline, IKI+ violet.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical; previously reported from Cuba, Colombia, Venezuela and Guyana.

Discussion. The conspicuous pseudostromata resemble those of Astrothelium degenerans and A. feei, but lack the internal yellow-orange pigment and are conspicuously flattened above. The amyloid ascospores also set A. infuscatulum apart.

New country record. Surinam: Brokopondo: Brownsberg, 1985, Aptroot 14865 (ABL).

Astrothelium inspersaeneum E. L. Lima et al.

In Lima et al. Bryologist 116: 327 (2013); type: Brazil, Pernambuco, Buíque, Catimbau National Park, on bark of tree, 900 m, 3 February 2012, Lima 394 (ISE!—holotype).

(Fig. 14G)

Thallus corticate, light olive-brown to yellowish, covered by orange pigment.

Ascomata trypethelioid, with apical ostioles, usually aggregated in groups or lines but not distinctly pseudostromatic, 0·2–0·4 mm diam., erumpent, covered by thallus and orange pigment except black ostiolar area. Hamathecium inspersed. Ascospores 8 per ascus, 3-septate, oblong-ellipsoid, 20–25×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K+ purple. TLC: an anthraquinone, probably parietin.

Distribution. Neotropical (previously reported only from Brazil).

Discussion. This species is most similar in aspect and internal structures to Astrothelium neogalbineum, but differs markedly by the inspersed hamathecium and the absence of lichexanthone.

New country record. Peru: San Martin: Tarapoto, Santesson & Thor P71:32 (S).

Astrothelium interjectum R. C. Harris

Acta Amazon. (Suppl.) 14: 61 (‘1984’) [1986]; type: Brazil, Amazonas, Serra Aracá, Samuels 838 (NY!—isotype).

(Fig. 24H & I)

Thallus light olive-green to greyish green, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 1–2(–3) mm diam., erumpent to prominent, whitish and darker ostiolar areas. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 25–30×6–11 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV+ yellow, K−, with lichexanthone.

Distribution. Pantropical; previously reported from Costa Rica and Brazil.

New country records. Colombia: Nariño: Tumaco, Estacion Forestal La Espriella, 1986, Sipman & Velosa 33041 (B).—Guyana: Upper Takutu: Kuyuwini Landing, 1992, Sipman 57017 (B).—Papua New Guinea: Madang: Gogol Valley, Tgubi logging site, 1992, Sipman 35965 (ABL, B); 11 km W of Brahman Mission, 1995, Aptroot 38706 (ABL).

Astrothelium intermedium Aptroot & Lücking

In Aptroot et al., Biblioth. Lichenol. 98: 46 (2008); type: Costa Rica, Puntarenas, Las Cruces Biological Station, Lücking s. n. (F!—holotype).

(Fig. 29G & H)

Thallus corticate, olive-green, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata diffusely pseudostromatic; pseudostromata 1–2 cm diam. (individual ascomata 0·3–0·5 mm diam.), erumpent, irregular to linear and in part confluent and anastomosing, covered by thallus except dark ostioles surrounded by a pale rim. Hamathecium clear. Ascospores 8 per ascus, 3-septate, 29–33×12–15 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Costa Rica).

Astrothelium irregulare (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816693

Bathelium irregulare Müll. Arg., Hedwigia 30: 234 (1891).—Laurera irregularis (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 504 (1922); type: Brazil, Theresopolis, Schenck 4643 (G!—holotype).

(Fig. 34F)

Thallus corticate, olive-green to brownish, uneven-verrucose.

Ascomata trypethelioid, with apical ostioles, solitary or rarely irregularly confluent, 0·6–1·0 mm diam., erumpent to prominent, covered by thallus. Hamathecium inspersed. Ascospores 8 per ascus, densely muriform, fusiform, 135–165×36–52 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Astrothelium isabellinum Eschw.

In Martius, Icon. Plant. Cryptog. 2: 20 (1828).—Astrothelium isabellinum Eschw. (1824), nom. nud.—Heufleria isabellina (Eschw.) Trevis., Flora 44: 23 (1861).—Cryptothelium isabellinum (Eschw.) Zahlbr., Catal. Lich. Univ. 1: 522 (1922); type: Brazil, s. col. (M—holotype, not seen).

Thallus corticate, olive-green, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata diffusely pseudostromatic; pseudostromata 1–2 mm diam., erumpent, covered by thallus, internally with red pigment. Hamathecium clear. Ascospores 2 per ascus, densely muriform, fusiform, 150–200×40–50 µm, without distinctly thickened median septum, hyaline, IKI–.

Chemistry. Thallus UV−, K−; pseudostromata UV−, externally K−, internally with K+ purple anthraquinone.

Distribution. Neotropical (Brazil).

Astrothelium keralense (Upreti & Ajay Singh) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816694

Laurera keralensis Upreti & Ajay Singh, Bull. Jard. Bot. Nat. Belg. 57: 374 (1987); type: India, Kerala, Quilon, Kundara, Singh & Ranjan 102951 (LWG—holotype, not seen).

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·6–1·0 mm diam., erumpent to prominent, hemispherical, covered by thallus. Hamathecium clear. Ascospores 8 per ascus, muriform, fusiform, 50–60×15–20 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (India and Thailand).

Astrothelium kunzei (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816695

Trypethelium kunzei Fée, Ann. Sci. Nat. 23: 445 (1831); type: Surinam, Kunze s. n. (G!—holotype).

(Fig. 15H & I)

Thallus corticate, olive-brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·5–1·0 mm broad, immersed, forming irregular, partly reticulate lines, covered by thallus and orange pigment except dark ostioles. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 20–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV+ red, K+ purple, with anthraquinone.

Distribution. Neotropical (Panama and Surinam).

Discussion. The epithet kunzei is taken up here for material in the Astrothelium aeneum aggregate that produce distinct, immersed, linear-reticulate pseudostromata. This distinction is supported by molecular data (Lücking et al. 2016 a ).

Astrothelium laevigatum Müll. Arg.

Flora 66: 245 (1883); type: Brazil, Apiahy, Puiggari 1689 (G!—holotype).

Astrothelium simplicatum Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 194 (1890); type: Brazil, Minas Gerais, Sítio, Vainio 1006 (TUR-Vain 30870!—holotype; BM!—isotype).

(Fig. 23K & L)

Thallus corticate, greenish grey (lichexanthone colour), smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, not pseudostromatic, 0·3–0·5 mm diam., erumpent, covered by thallus but upper part whitish contrasting with the dark ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 20–26×7–9 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone).

Distribution. Neotropical (Brazil).

Astrothelium leioplacum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816696

Clathroporina leioplaca Müll. Arg., Revue Mycol. 10: 182 (1888); type: Paraguay, Balansa s. n. (G!—holotype).

Clathroporina irregularis Müll. Arg., Revue Mycol. 10: 182 (1888); type: Paraguay, Balansa 231 (G!—lectotype, designated here).

(Fig. 39H)

Thallus corticate, olive-grey, smooth to uneven.

Ascomata pleurothelioid, with separate ostioles, 0·4–0·6 mm diam., erumpent, completely covered by thallus. Hamathecium clear. Ascospores 8 per ascus, densely muriform, fusiform, 33–52×16–22 µm, without thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Paraguay).

Astrothelium leucoconicum Nyl.

Flora 52: 126 (1869); type: Brazil, Rio de Janeiro, Glaziou s. n. (H-Nyl 118!—holotype).

(Fig. 24K & L)

Thallus corticate, light olive-green to yellowish, uneven to shallowly bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, not pseudostromatic, 0·4–0·7 mm diam., erumpent, covered by thallus but upper part whitish contrasting with the dark ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 50–72×20–26 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone); ascomata UV−, K−.

Distribution. Neotropical (previously reported from Brazil).

New country record. Venezuela: Bolivar: Cerro Guaiquinima, 1500 m, 1990, Sipman 27215 (B).

Astrothelium leucothelium Nyl.

Ann. Sci. Nat., Bot., sér. 5 7: 348 (1867); type: Colombia, Rio Negro, Lindig s. n. (H-Nyl 117!—holotype; S!—isotype).

(Fig. 23G)

Thallus corticate, greenish grey (lichexanthone colour), becoming brownish yellow, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata solitary to confluent or diffusely pseudostromatic, 0·4–0·6 mm diam., erumpent, covered by thallus but upper part whitish, with dark ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 3(–5)-septate, fusiform, 32–52×10–17 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−. TLC: lichexanthone.

Distribution. Neotropical (previously reported from Costa Rica, Colombia and Brazil).

New country records. Venezuela: Merida: Moro Negro, 1980, López Figueras & Rodríguez 22888 (ABL, MERF).—French Guiana: Saül, 1988, Sipman 31769, 31757 (ABL, B).

Astrothelium lugescens (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816697

Verrucaria lugescens Nyl., Flora 69: 177 (1886).—Anthracothecium lugescens (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 463 (1922).—Campylothelium lugescens (Nyl.) Upreti & Ajay Singh, Bull. Soc. Bot. Fr., Let. Bot. 134: 293 (1987); type: São Tomé and Príncipe, Henriques s. n. (H-Nyl 1067!—holotype).

(Fig. 38I)

Thallus corticate, olive-green, smooth to uneven.

Ascomata pleurothelioid, with separate ostioles, 0·6–1·0 mm diam., erumpent, covered by thallus. Hamathecium clear. Ascospores 1 per ascus, densely muriform, fusiform, 210–230×45–70 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. African palaeotropical (São Tomé and Príncipe).

Astrothelium luridum (Zahlbr.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816698

Trypethelium luridum Zahlbr., Ann. Mycol. 14: 47 (1916); type: Japan, Thushima, Faurie 3744 (W—holotype, not seen).

Trypethelium scoria f. endochraceum Nyl, Lich. Jap.: 115 (1890); type: Malaysia, Labuan, Almquist, 1897 (S!—isotype).

Trypethelium endosulphureum Makhija & Patw., J. Hattori Bot. Lab. 73: 197 (1993); type: India, Meghalaya, Mohmtheid-Cherapunji road, 20 October 1977, Patwardhan & Nagarkar AMH 77.905 (ABL!—isotype).

(Fig. 17F–H)

Thallus corticate, olive-brown to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·8–1·5 mm broad, erumpent to prominent, covered by thallus except for dark ostiolar area, internally with yellow pigment. Hamathecium clear. Ascospores 8 per ascus, 7–11-septate, fusiform, 55–88×13–28 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata U−, internally K+ red, with anthraquinone.

Distribution. Asia, extending into temperate regions (India and Japan).

Astrothelium macrocarpum (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816699

Porina macrocarpa Fée, Essai Crypt. Écorc.: 81 (1824); non Riddle 1920.—Porophora macrocarpa (Fée) Spreng., Syst. Veget. 4(1): 241 (1827).—Pyrenodium macrocarpum (Fée) Fée, Essai Crypt. Écorc. Suppl.: 69 (1837).—Pyrenula macrocarpa (Fée) Massal., Ricerch. Auton. Lich.: 164 (1852).—Astrothelium hypoxylon var varmacrocarpum (Fée) Nyl., Mém. Soc. Sci. Nat. Cherbourg 5: 141 (1857).—Astrothelium sulphureum var. macrocarpum (Fée) Nyl., in Hue, Nouv. Archiv. du Muséum, sér. 3 4: 132 (1892); type: South America, “Cort. cinchonae” (M!—isotype).

Astrothelium galbineum Kremp., Nuovo Giorn. Bot. Ital. 7: 58 (1875); type: Sarawak, Beccari 23 (M!—lectotype; Harris, Acta Amazon. (Suppl.) 14: 60 (‘1984’) [1986]).

Astrothelium ochrothelizum Müll. Arg., Bot. Jahrb. Syst. 6: 382 (1885).—Trypethelium ochrothelizum Nyl., Flora 59: 365 (1876), nom. nud.—Trypethelium ochrothelizum (Müll. Arg.) Nyl., Sert. Lich. Trop. Labuan et Singap.: 26 (1891); type: Cuba, Wright s. n. (G!—holotype; BM!—isotype; Müller, Verr. Cub. 144).

Astrothelium conicum var. pallidum Müll. Arg., Bot. Jahrb. Syst. 6: 382 (1885); type: Cuba, Wright s. n. (G!—lectotype, designated here; BM!—isolectotype; Müller, Verr. Cub. 605).

Astrothelium ochrothelioides Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 194 (1890); type: Brazil, Minar Gerais, Lafayette, Vainio s. n. (TUR-Vain 30868!—lectotype, designated here; Vainio, Lich. Bras. Exs. 310).

Trypethelium discolor Müll. Arg., Hedwigia 34: 35 (1895); type: Brazil, Minas Gerais, Serra do Ouro Preto, Ule 298 (G!—holotype).

(Fig. 22G–K)

Thallus corticate, olive-green to yellowish, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 0·5–1·0 mm diam., erumpent to prominent, conical with flattened top, covered by thallus with lateral part whitish and upper part with yellow-orange pigment and dark ostiole. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 21–28×7–11 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone); pseudostromata with yellow-orange K+ purple, UV+ red anthraquinone, probably parietin.

Discussion. The rather well-known name Astrothelium galbineum has to be replaced by the older epithet macrocarpum, which was in use for a century but then surprisingly went out of use. The synonymy of A. ochrothelioides is based on the original description of the ascospores; Harris (1986: 59) reported that no spores remain. Typical A. macrocarpum has conical pseudostromata in which only the upper part is covered by pigment, whereas the types of A. ochrothelioides and Trypethelium discolor have wart-shaped pseudostromata completely covered by pigment; thus, more than one species might be contained here.

Distribution. Pantropical; previously reported from the USA, Cuba, Costa Rica, Panama, Venezuela, Guyana, French Guiana, South Africa, India, Sri Lanka, Singapore, Thailand, Borneo, Papua New Guinea and Australia.

New country records. Colombia: Amazonas: Araracuara, opposite airstrip, 350 m, 1988, Sipman & Duivenvoorden 27863 (B).—Gabon: Nzé, 2006, Ertz 9719 (BR).

Astrothelium macrosporum (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816700

Trypethelium macrosporum Makhija & Patw., J. Hattori Bot. Lab. 73: 200 (1993); type: India, Meghalaya, Cherapunji Road, 27 October 1977, Patwardhan & Nagarkar AMH 77.826 (ABL!—isotype).

(Figs 21F & 33J)

Thallus corticate, olive-green to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·7–1·0 mm diam., erumpent, covered by thallus except dark ostiolar area surrounded by irregular whitish rim. Hamathecium finely inspersed. Ascospores 8 per ascus, 9–13-septate, occasionally with a longitudinal septum, fusiform, 120–195×17–22 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (India).

Astrothelium marcidum (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816701

Pyrenula marcida Fée, Essai Crypt. Écorc.: 77 (1824).—Verrucaria marcida Spreng. (1827) nom. nud.—Trypethelium marcidum (Fée) Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 30(3): 11 (1888); type: South America “ad corticem Cinchonarum”, “b” (G!—holotype).

(Fig. 29F)

Thallus corticate, olive-brown, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic, pseudostromata 0·7–1·0 mm diam., immersed-erumpent, covered by thallus or yellowish white layer. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 35–45×14–16 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical.

Discussion. The epithet marcidum was suggested to replace the epithet floridanum by Aptroot et al. (2008), as both agree in thallus morphology, ascospore type, and lack of secondary substances. The type material of Pyrenula marcida is in very bad shape, with almost all ascomata abraded; however, re-examination revealed that the ascoma organization is astrothelioid, not trypethelioid as in Astrothelium floridanum, since several remaining ascoma bases show a pyriform shape with the tip pointing centrally.

Astrothelium megaleium (Kremp.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 8166702

Trypethelium megaleium Kremp., Nuovo Giorn. Botan. Ital. 21(3): 13 (1875).—Bathelium megaleium (Kremp.) Müll. Arg., Linnaea 63: 45 (1880).—Laurera megaleia (Kremp.) Zahlbr., Catal. Lich. Univ. 1: 505 (1922); type: Malaysia, Sarawak, Beccari s. n. (M!—holotype).

?Trypethelium oligosporum Mont. & Bosch, in Mont., Sylloge Gener. Spec. Cryptog.: 374 (1865).—Bathelium oligosporum (Mont. & Bosch) Trevis., Flora 44: 21 (1861).—Laurera oligospora (Mont. & Bosch) Zahlbr., Catal. Lich. Univ. 1: 505 (1922); type: Indonesia, Java, Junghuhn s. n. (L—holotype lost).

(Fig. 33I)

Thallus corticate, olive-brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·8–1·5 mm diam., erumpent, mostly exposed, dark brown. Hamathecium inspersed. Ascospores 2 per ascus, densely muriform, fusiform, 210–225×32–35 µm, with distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (Malaysia: Sarawak, and Indonesia).

Discussion. The synonymy of Trypethelium oligosporum is uncertain, as the type is lost, and therefore this older epithet is not taken up.

Astrothelium megalophthalmum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816703

Trypethelium megalophthalmum Müll. Arg., Hedwigia 34: 35 (1895); type: Brazil, Santa Catarina, Serra Geral, Ule 291 (G!—holotype; NY—isotype).

(Fig. 22A & B)

Thallus corticate, olive-green, verrucose-bullate.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic; pseudostromata 1–3(–5) mm diam., immersed-erumpent, with upper portion of ascomata exposed, blackish brown. Hamathecium clear. Ascospores 8 per ascus, 3–7-septate, fusiform, 85–120×20–32 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Astrothelium megalostomum (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816704

Heufleria megalostoma Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 193 (1890).—Cryptothelium megalostomum (Vain.) Zahlbr., Catal. Lich. Univ. 1: 522 (1922).—Campylothelium megalostomum (Vain.) Aptroot, Fungal Diversity 9: 29 (2002); type: Brazil, Minas Gerais, Serra do Caraça, Vainio s. n. (TUR-Vain 303838!—holotype; Vainio, Lich. Bras. Exs. 1587).

(Fig. 36E)

Thallus corticate, light olive-green, smooth to rough.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 0·8–1·3 mm diam., erumpent, covered by thallus except dark ostiolar area. Hamathecium inspersed. Ascospores 2 per ascus, densely muriform, fusiform, 170–220×50–64 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata UV+ yellow, with lichexanthone, K−.

Distribution. Neotropical (Brazil).

Astrothelium megaspermum (Mont.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816705

Trypethelium megaspermum Mont., Ann. Sci. Nat., Bot., sér. 2 19: 68 (1843).—Bathelium megaspermum (Mont.) Trevis., Spighe e Paglie: 20 (1861).—Laurera megasperma (Mont.) Riddle, Bull. Torrey Bot. Club 44: 323 (1917); type: French Guiana, Leprieur 603 (FH-TAYL 11!—isotype).

Trypethelium ostendatum Kremp., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 5: 398 (1874).—Bathelium ostendatum (Kremp.) Müll. Arg., Linnaea 63: 45 (1880).—Laurera ostendata (Kremp.) Zahlbr., Catal. Lich. Univ. 1: 506 (1922); type: Brazil, Minas Gerais, Serra d’Estrella, Warming 79 (M!—holotype).

Verrucaria euthelia Nyl., Flora 69: 177 (1886); type: São Tomé and Príncipe, Moller s. n. (H-Nyl 1064! —holotype).

Thelenella fulva Vain., Catal. Welwitsch Afric. Plants 2: 451 (1901).—Clathroporina fulva (Vain.) Zahlbr., Catal. Lich. Univ. 1: 418 (1922).—Polyblastiopsis fulva (Vain.) C. W. Dodge, Ann. Missouri Bot. Gard. 40: 275 (1953); type: Angola, Golungo Alto, Mata Quisuculo, Welwitsch 230 (TUR-Vain 31066!—holotype; BM!—isotype).

Julella zenkeriana P. Henn., Bot. Jahrb. Syst. 38: 127 (1905); type: Cameroon, Bipinde, Zenker 1980 (B!—holotype).

Polyblastiopsis pertusarina Zahlbr., Feddes Repert. 31: 199 (1933); type: Taiwan, Mount Arisan, Toroyen, Asahina 299 (W—holotype, not seen).

Campylothelium nitidum Zahlbr., Ann. Mycol. 33: 39 (1935) non Müll. Arg. (1891).—Campylothelium zahlbruckneri R. G. Werner, Bull. Soc. Sci. Nat. Maroc. 24: 130 (1944); type: USA, Florida, Sanford, Rapp 84 (W—holotype, not seen).

Clathroporina diphloea Zahlbr., Ann. Mycol. 33: 37 (1935); type: USA, Florida, Rapp (W—holotype, not seen).

Laurera megasperma f. immersa Letr.-Gal., Rev. Bryol. Lichénol. 26: 234 (1957); type: Sri Lanka, Petsch (BM!—holotype).

Laurera megasperma f. conica Letr.-Gal., Rev. Bryol. Lichénol. 27: 67 (1958); type: Ivory Coast, Mont Orombe Boka, Dimbokro, 20 August 1954, Santesson 10724o (UPS—holotype, not seen).

(Fig. 34H–L)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·8–1·5 mm diam., prominent, hemispherical with flattened top, covered by thallus except for dark ostiolar area surrounded by whitish rim. Hamathecium inspersed. Ascospores generally 4 per ascus, densely muriform, oblong-fusiform, 140–220×30–75 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Pantropical; previously reported from the USA, Costa Rica, Cuba, Jamaica, Puerto Rico, French Guiana, Brazil, Argentina, Ivory Coast, Angola, Cameroon, India, Sri Lanka, Thailand, Vietnam, Taiwan, New Caledonia and Papua New Guinea.

New country records. Colombia: Nariño: Tumaco, Estacion Forestal La Espriella, 1986, Sipman & Velosa 33054 (B).—Guyana: Upper Takutu: Kuyuwini Landing, 1992, Sipman 58222 (B).—Ecuador: Napo: 5 km N of Sta. Rosa, 400 m, 1982, Aptroot & Hensen 10611, 10612 (ABL).—Peru: Maynas: Iquitos, 1980, Schumm & Pudor 11952 (hb. Schumm).—Gabon: Nzé, 2006, Ertz 9725 (BR).—DR Congo: Mongala: Lisala, 2009, Ertz 14181 (BR).—Malaysia: Pahang: Fraser’s Hill, 1989, Sipman et al. 45022 (B).—China: Yunnan: Xishuangbanna, Menglun, 2002, Aptroot 57355 (ABL).—Indonesia: Sulawesi: Tondano, 1000 m, 1988, Hensen s. n. (ABL).

Astrothelium meiophorum (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816732

Trypethelium annulare var. meiophorum Nyl., Bull. Soc. Linn. Normand., sér. 2 2: 132 (1868).—Trypethelium meiophorum (Nyl.) Müll. Arg., Hedwigia 31: 286 (1892); type: New Caledonia, Pancher s. n. (G!—lectotype, designated here; BM!—isolectotype).

Trypethelium flavoalbum Makhija & Patw., J. Hattori Bot. Lab. 73: 197 (1993); type: India, Megahalya, near Sohararim, 20 October 1977, Patwardhan & Nagarkar AMH 86.697 (ABL!—isotype).

Trypethelium parvicarpum Makhija & Patw., J. Hattori Bot. Lab. 73: 205 (1993); type: India, Nicobar Islands, Kamorta, Daring, 18 January 1987, Nagarkar & Patwardhan AMH 87.389 (ABL!—isotype).

(Fig. 19C & D)

Thallus corticate, light olive-green to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, irregularly grouped to diffusely pseudostromatic; pseudostromata 0·5–0·7 mm broad, erumpent, covered by thallus except for broad, dark ostiolar area. Hamathecium clear. Ascospores (2–)8 per ascus, fusiform-ellipsoid, 3-septate, 20–25×7–8 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (previously reported from India and New Caledonia).

Discussion. Unfortunately the type material of Trypethelium annulare var. meiophorum is extremely small and hence the new synonyms listed here are somewhat tentative, although morphologically they seem to agree well, particularly the type of T. parvicarpum. We were unable to confirm the statement of 2-spored asci given in the protologue of Astrothelium meiophorum, since only two ascomata are left in the type, but we believe that this observation is based on an ascus with most ascospores already expelled. The two synonyms listed regularly have 8-spored asci. Additional material found under this name from New Caledonia is A. scoria. It should also be mentioned that the ascospores in the type specimen of Trypethelium parvicarpum are c. 23×7 µm, larger than given in the protologue of that taxon.

New country record. China: Yunnan: Xishuangbanna, 2002, Aptroot 57350 (ABL).

Astrothelium meristosporoides (P. M. McCarthy & Vongshew.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816706

Laurera meristosporoides P. M. McCarthy & Vongshew. in Vongshewarat et al., Mycotaxon 70: 228 (1999); type: Thailand, Phisanulok Prov., Phu Hin Rong Kla National Park, Vongshewarat 173 (CANB—holotype, not seen).

(Fig. 31E)

Thallus corticate, light greenish yellow, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, prominent, whitish and black ostiolar area, 0·7–1·5 mm diam. Hamathecium inspersed. Ascospores 8 per ascus, muriform, fusiform, 68–120×18–24 µm, with somewhat thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−, with lichexanthone or 1,8-dihydroxy-3,6-dimethoxyxanthone.

Distribution. Palaeotropical (previously reported only from Thailand).

New country record. Papua New Guinea: Central: Varirata National Park, 800 m, 1987, Aptroot 19135 (ABL).

Astrothelium meristosporum (Mont. & Bosch) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816707

Trypethelium meristosporum Mont. & Bosch, in Junghuhn, Pl. Jungh. 4: 487 (1855).—Meristosporum javanicum A. Massal., Atti I. R. Istitut. Veneto, Ser. 3 5: 328 (1860).—Bathelium meristosporum (Mont. & Bosch) Trevis., Flora 44: 21 (1861).—Melanotheca javanica (A. Massal.) Zahlbr., in Engler & Prantl, Natürl. Pflanzenfamil. 1(1*): 70 (1903).—Laurera meristospora (Mont. & Bosch) Zahlbr., Catal. Lich. Univ. 1: 505 (1922); type: Indonesia, Java, Junghuhn (L!—isotype).

Bathelium chrysocarpum Müll. Arg., Hedwigia 30: 54 (1891).—Laurera chrysocarpa (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 503 (1922); type: Australia, Queensland, Bellenden Ker, Bailey 540 (G! —holotype; BRI!—isotype).

Verrucaria anoptella Stirt., Proceed. Philosoph. Soc. Glasgow 13: 190 (1881).—Clathroporina anoptella (Stirt.) Zahlbr., Catal. Lich. Univ. 1: 415 (1922).—Polyblastiopsis anoptella (Stirt.) Ajay Singh, Nova Hedwigia 36: 238 (1982); type: India, Assam, Watt s. n. (BM!—holotype).

Thelenella interrupta Vain., Hedwigia 46: 180 (1907).—Clathroporina interrupta (Vain.) Zahlbr., Catal. Lich. Univ. 1: 418 (1922); type: Thailand, Koh Chang Island, Lem Dam, Schmidt 17 (TUR-Vain 31065—holotype, not seen).

Laurera columellata Makhija & Patw., Mycotaxon 31: 574 (1988); type: India, Karnataka, S. Kanara, Hiriyadaka, Udupi-Hebri Road, 22 February 1978, Patwardhan AMH 78.59 (ABL!—isotype).

Laurera andamanica D. D. Awasthi, Biblioth. Lichenol. 40: 3 (1991).—Laurera indica Makhija & Patw., Mycotaxon 31: 578 (1988) non Upreti & Ajay Singh (1987); type: India, Andaman Islands, South Andaman, Baratang Islands, Bishnu Nala, 22 February 1985, Patwardhan & Sethy AMH 85.656 (ABL!—isotype).

(Fig. 31F–H)

Thallus corticate, light olive-green, uneven to verrucose.

Ascomata trypethelioid, with apical ostioles, 1·0–1·5 mm diam., prominent, hemispherical with flattened top, covered by thallus except whitish to greyish ostiolar area, 0·7–2·0 mm diam. Hamathecium hyaline or slightly yellowish, inspersed. Ascospores (4–)8 per ascus, muriform, oblong-fusiform, 120–220×25–40 µm, constricted at the markedly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV+ yellow, K−, with lichexanthone or 1,8-dihydroxy-3,6-dimethoxyxanthone.

Distribution. Palaeotropical (previously reported from India, Sri Lanka, Thailand, Indonesia, Papua New Guinea, Australia); the specimen reported here from French Guiana is atypical.

Discussion. As can be expected for a species with such large ascospores, there is considerable variation in size, even within a single ascoma.

New country records. French Guiana: Montsinery, 20 km W of Cayenne, along forest track called Risque tout, 50 m, 1985, Aptroot 15125 (ABL, B).—Malaysia: Belum, Halong Trail, 1994, Nätebusch s. n. (hb. Kalb).—Solomon Islands: Kolombangara, 1965, Hill 10377 (BM).

Astrothelium neogalbineum (R. C. Harris) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816708

Trypethelium neogalbineum R. C. Harris, Acta Amazon. (Suppl.) 14: 74 (‘1984’) [1986]; type: Brazil, Pará, Santarém, Spruce 259 (BM!—isotype).

(Fig. 12A & B)

Thallus corticate, olive-green to greyish green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·5–1·0 mm broad, immersed-erumpent, irregular to elongate or becoming reticulate, dark brown with orange pigment, especially laterally. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 18–26×7–8 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, with lichexanthone; pseudostromata UV+ red, K+ purple, with anthraquinone.

Distribution. Neotropical (previously reported from Costa Rica and Brazil).

New country record. Guyana: East Demarara: along Linden Highway, Soesdyke, Sipman 40277 (B).

Astrothelium nigratum (Müll. Arg.) Aptroot & Lücking comb. et stat. nov.

MycoBank No.: MB 816709

Astrothelium minus var. nigratum Müll. Arg., Bot. Jahrb. Syst. 6: 382 (1885); type: Cuba, Wright s. n. (G!—holotype; FH-TUCK 3982—isotype, not seen; Müller, Verr. Cub. 638).

(Fig. 19B)

Thallus corticate, olive-brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic, 0·2–0·3 mm diam., immersed-erumpent, exposed and brown-black. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 20–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported only from Cuba).

New country records. Puerto Rico: Mayagüez: Reserva Forestal Maricao, 1989, A. & M. Aptroot 24961 (ABL).—Guyana: Rupununi: Kuyuwini Landing, 1991, Jansen-Jacobs et al. 2573 (ABL).

Astrothelium nigricans Malme

Astrothelium nigricans Malme, Ark. Bot. 19(1): 13 (1924); type: Brazil, Matto Grosso, Cuyabá, Malme 2003 (S!—holotype).

(Fig. 28F)

Thallus corticate, olive-green to yellowish, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata diffusely pseudostromatic, formed on irregularly linear to reticulate, whitish thallus portions, 0·6–1·0 mm diam., prominent, wart-shaped, completely exposed and black, covered. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 21–27×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Astrothelium nigrorufum (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816710

Trypethelium nigrorufum Makhija & Patw., J. Hattori Bot. Lab. 73: 203 (1993); type: India, Andaman Islands, Baratang, Baludera, 23 February 1985, Patwardhan & Sethy AMH 85.845 (ABL!—isotype).

(Fig. 17A)

Thallus corticate, olive-brown to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 1–2 mm broad, immersed-erumpent, exposed and brown-black, internally with yellow-orange, K+ purple pigment. Hamathecium clear. Ascospores 8 per ascus, 9–13-septate, fusiform, 29–35(–46)×7–9 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV−, internally K+ purple, with anthraquinone.

Distribution. Eastern palaeotropical (India).

Astrothelium nitidiusculum (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816711

Verrucaria nitidiuscula Nyl., Ann. Sci. Nat. Bot. sér. 4 20: 252 (1863).—Pseudopyrenula nitidiuscula (Nyl.) Müll. Arg., Flora 66: 248 (1883).—Trypethelium nitidiusculum (Nyl.) R. C. Harris, Acta Amazon. (Suppl.) 14: 75 (‘1984’) [1986]; type: Colombia, Villeta, Lindig 2829 (M!, BR—isotypes).

Pseudopyrenula neglecta Müll. Arg., Flora 68: 332 (1885); type: French Guiana, Leprieur 479 (G!—holotype).

Trypethelium hebetum Makhija & Patw., J. Hattori Bot. Lab. 73: 198 (1993); type: India, Andaman Islands, South Andaman, Near Wright Myo, Shol Bay, 20 December 1986, Patwardhan & Nagarkar AMH 77.866 (ABL!—isotype).

(Fig. 19F–L)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to confluent but not distinctly pseudostromatic, 0·2–0·3 mm diam., erumpent, covered by thallus but upper portion whitish (or a white rim only) with dark ostiolar area. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 15–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical, reported from, for example, Colombia, Ecuador, Bolivia, Guyana, French Guiana, Brazil, India, and Papua New Guinea. The species has also been reported from the USA, Mexico, Bermudas, Bahamas, Cuba, Jamaica, Dominican Republic, Trinidad, Costa Rica, El Salvador, Isla del Coco, Guyana, French Guiana, Colombia, Venezuela, Galapagos, Brazil, Bolivia, Argentina, South Africa, Seychelles, India, Sri Lanka, Vietnam, Thailand, Taiwan, Hong Kong, Japan, Papua New Guinea and Australia, but these reports correspond to a much broader species concept including, for example, the now separated, more common taxa A. bicolor and A. scoria. Therefore, below are some additional records from countries from which the species had been reported previously in a wider sense.

Discussion. Astrothelium nitidiusculum is restricted here to include specimens with solitary to confluent, but not pseudostromatic ascomata. Even when dense or confluent, the ascomata can clearly be individually discerned.

Confirmed additional country records. Ecuador: Morona-Santiago: Gualaquiza, 1982, Aptroot & Hensen 10454 (ABL).—Guyana: Upper Takutu: Kuyuwini Landing, 1992, Sipman 58140 (B).—Papua New Guinea: Madang: Balek Wildlife Reserve, 1992, Aptroot 30891 (ABL).

Astrothelium nitidulum Weerakoon & Aptroot

Cryptog., Mycol. 35: 58 (2014); type: Sri Lanka, Central Province, Coolbone tea estate Matale, Cloonan 083/53, 1982 (F!—holotype).

(Fig. 21E)

Thallus corticate, olive-brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to confluent and becoming diffusely pseudostromatic; pseudostromata 0·9–1·9 mm diam. (individual ascomata 0·6–0·9 mm diam.), erumpent to prominent, covered by thallus except for black ostiole with whitish centre. Hamathecium inspersed. Ascospores 8 per ascus, 7-septate, fusiform, 40–49×12–15 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−. TLC: no substances detected.

Distribution: Eastern palaeotropical (Sri Lanka).

Astrothelium obscurum Müll. Arg.

Flora 66: 244 (1883); type: Brazil, Apiahy, Puiggari 2255 (G!—holotype).

(Fig. 28J)

Thallus corticate, olive-brown, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata largely confluent but not distinctly pseudostromatic, 0·3–0·4 mm diam., erumpent, applanately conical, exposed and dark brown. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 20–25×7–9 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Astrothelium ocellatum Malme

Ark. Bot. 19(1): 14 (1924); type: Brazil, Matto Grosso, Santa Anna da Chapada, Malme 2411 (S!—holotype).

(Fig. 27D & E)

Thallus corticate, olive-green, smooth to uneven or shallowly bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata solitary to confluent but not pseudostromatic, 1·0–1·8 mm diam., erumpent, covered by thallus but papilliform ostiolar area ochraceous brown, with yellow-orange pigment and surrounded by an irregular whitish rim. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 55–65×15–25 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−, without lichexanthone; pseudostromata with yellow-orange, K+ purple, UV+ red anthraquinone.

Distribution. Pantropical (previously reported from Guyana, Brazil and Papua New Guinea).

New country records. Venezuela: Amazonas: Alto Orinoco, Surumoni, 1998, Hafellner & Komposch 15-13 (GZU).—French Guiana: Saül, 1988, Sipman 31772 (B).

Astrothelium ochrothelium (Nyl.) Müll. Arg.

Flora 68: 255 (1885).—Trypethelium ochrothelium Nyl., Acta Soc. Sci. Fenn 7: 494 (1863); type: Colombia, Villeta, Lindig 2823 (H-Nyl 7574!—lectotype; Harris, Acta Amazon. (Suppl.) 14: 62 (‘1984’) [1986]; S!, BR—isolectotypes).

(Fig. 23A & B)

Thallus corticate, olive-green to yellowish or brownish, uneven to shallowly verrucose.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 0·8–1·5 mm broad, erumpent, irregular, often confluent, covered by thallus and upper portion with yellow-orange pigment. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, (37–)43–45×15–23 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone; pseudostromata UV−, K+ purple, with yellow-orange anthraquinone.

Distribution. Neotropical (previously reported from Cuba, Guyana, Venezuela, Colombia and Brazil).

New country record. French Guiana: Kourou, 1958, Degelius s. n. (ABL, UPS).

Astrothelium octosporoides Aptroot & Lücking nom. nov.

MycoBank No.: MB 816712

Bathelium octosporum Zahlbr., Sitzungsber. K. Akad. Wiss. Wien, math.-naturw. Klasse 111(1): 372 (1902).—Laurera octospora (Zahlbr.) Zahlbr., Catal. Lich. Univ. 1: 505 (1922) non Astrothelium octosporum (Vain.) Aptroot & Lücking (see below); type: Brazil, Theresiopolis, von Höhnel 150 (W—holotype, not seen).

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·8–1·5 mm diam., erumpent to prominent, hemispherical, covered by thallus. Hamathecium clear. Ascospores 8 per ascus, densely muriform, fusiform, 150–190×40–45 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Astrothelium octosporum (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816713

Heufleria octospora Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 192 (1890).—Cryptothelium octosporum (Vain.) Zahlbr., Catal. Lich. Univ. 1: 522 (1922); type: Brazil, Minas Gerais, Sitio, Vainio s. n. (TUR-Vain 30839!—holotype; BM—isotype; Vainio, Lich. Bras. Exs. 1031).

(Fig. 37D)

Thallus corticate, olive-green to brownish, uneven to shallowly bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 0·8–1·7 mm diam., immersed to erumpent, covered by thallus except dark ostiolar area surrounded by thin whitish rim. Hamathecium clear. Ascospores 8 per ascus, densely muriform, narrowly fusiform, 110–135×19–23 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−. TLC: lichexanthone.

Distribution. Neotropical (Brazil).

Astrothelium oligocarpum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816714

Trypethelium oligocarpum Müll. Arg., Nuovo Giorn. Bot. Ital. 23: 402 (1891); type: Australia, Queensland, Brisbane, Bailey 465 p.p. (G!—lectotype, designated here).

(Fig. 28K)

Thallus corticate, olive-green to yellowish, shallowly bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata grouped but not distinctly pseudostromatic, 0·3–0·4 mm diam., prominent, basally covered by thallus but upper part blackish. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 20–27×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (Australia).

Astrothelium olivaceofuscum (Zenker) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816715

Trypethelium olivaceofuscum Zenker, in Goebel & Kunze, Pharmazeut. Waarenkunde 1(3): 190 (1827); type: South America (L!—holotype).

(Fig. 22C & D)

Thallus corticate, brownish to light greyish green or yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to often irregularly confluent but not distinctly pseudostromatic, 0·6–1·2 mm diam., erumpent, covered by thallus but protruding ostiole dark. Hamathecium clear. Ascospores 8 per ascus, 9–17-septate, fusiform, 85–100×8–16 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Costa Rica).

New country records. Guyana: Potaro-Siparuni Region: surroundings of Paramakatoi Village, 800 m, 1996, Sipman 41275 p.p. (B).—Brazil: Alagoas: Pilar, 2001, Cáceres & Lücking s. n. (ABL, URM).

Astrothelium papillosum (P. M. McCarthy) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816716

Laurera papillosa P. M. McCarthy, Lichenologist 27: 311 (1995); type: Papua New Guinea, Central Prov., 40 km NE of Port Moresby, near Dabamura, on Owers Corner Rd, 580 m, 10 February 1981, Streimann & Naomi 14908 (B!—isotype).

(Fig. 34D)

Thallus corticate, green, distinctly verrucose-papillose.

Ascomata trypethelioid to pleurothelioid, with apical or eccentric but then separate ostioles, 0·6–1·0 mm diam., immersed and covered by thallus except for brownish black ostioles. Hamathecium orange, inspersed. Ascospores 8 per ascus, muriform, fusiform-ellipsoid, 130–195×26–34 µm, constricted at the markedly thickened median septum, hyaline, IKI−.

Chemistry. Thallus UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (Papua New Guinea).

Astrothelium papulosum (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816717

Verrucaria papulosa Nyl., Ann. Sci. Nat. Bot., sér. 5 7: 345 (1867).—Pseudopyrenula papulosa (Nyl.) Müll. Arg., Flora 66: 248 (1883).—Trypethelium papulosum (Nyl.) Makhija & Patw., Int. J. Mycol. Lichenol. 5: 242 (1992); type: Colombia, Pie de Cuesta, Lindig 96 (H-Nyl 1227!—lectotype; BR—isolectotype; Makhija & Patwardhan, Int. J. Mycol. Lichenol. 5: 242, 1992).

(Fig. 20E)

Thallus corticate, yellowish brown, uneven to verrucose-papillose.

Ascomata trypethelioid, with apical ostioles, aggregate in diffuse pseudostromata; pseudostromata 1–3(–5) mm diam. (individual ascomata 0·2–0·4 mm diam.), immersed to erumpent, covered by thallus except for dark ostioles. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 26–38×7–9 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Costa Rica and Colombia).

New country record. Brazil: Amapá: Floresta Nacional do Amapá, 2015, Cáceres & Aptroot 27267 (ABL, ISE).

Astrothelium peranceps (Kremp.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816718

Trypethelium peranceps Kremp., Nuovo Giorn. Bot. Ital. 7: 55 (1875); type: Malaysia, Sarawak, Beccari 112 (M!—lectotype; Harris, Acta Amazon. (Suppl.) 14: 60 (‘1984’) [1986]).

(Fig. 28E)

Thallus corticate, brownish, thin, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 0·3–0·4 mm diam., prominent, conical to wart-shaped, covered by thallus but upper portion often exposed and dark brown. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 21–27×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (Malaysia: Sarawak).

Discussion. This name has been reinstated as a separate species in the Astrothelium crassum complex. This complex is still poorly understood, but the considerable morphological differences between specimens that agree in ascoma organization, ascospore type, and lack of secondary substances suggests the existence of several taxa. Within this complex, A. peranceps is distinguished by its thin thallus and widely dispersed, prominent and conical to wart-shaped ascomata.

Astrothelium phaeothelium (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816719

Trypethelium phaeothelium Nyl., Ann. Sci. Nat., Bot., sér. 5 7: 347 (1867); type: Colombia, Rio Magdalena, Lindig 31 (H-Nyl 271!—holotype; BM!, BR, S!—isotypes).

(Fig. 20J)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, irregularly confluent to pseudostromatic; pseudostromata 1–2 mm broad (individual ascomata 0·8–1·2 mm broad), prominent to sessile, blackish brown, with slightly protruding, black ostioles. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 75–90×17–23 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Colombia).

Discussion. This is another species morphologically similar to Bathelium but it has been included in Astrothelium due to the lack of internal pigment in the pseudostromata and the astrothelioid ascospores.

Astrothelium phlyctaena (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB816720

Trypethelium phlyctaena Fée, Essai Crypt. Écorc.: 68 (1824).—Trypethelium scoria var. phlyctaena (Fée) Trevis., Flora 44: 19 (1861); type: Saint Lucia, s. col. (G!—lectotype, designated here; BM!—isolectotype).

Verrucaria catervaria Fée, Essai Crypt. Écorc.: 90 (1824).—Pyrenula catervaria (Fée) Massal., Framment. Lich: 25 (1855).—Spermatodium catervarium (Fée) Trevis., Conspect. Verruc.: 10 (1860).—Pseudopyrenula catervaria (Fée) Müll. Arg., Flora 66: 248 (1883).—Trypethelium catervarium (Fée) Tuck., Genera Lich.: 260 (1872); type: South America, “corticem cinchonarum” (G!—lectotype; Harris, Acta Amazon. (Suppl.) 14: 64 (‘1984’) [1986]).

Verrucaria decolorata Fée, Essai Crypt. Écorc.: 91 (1824); type: South America (G!—holotype; L!—isotype).

Verrucaria tessella Pers., in Gaudichaud, Voy. Uranie: 183 (1827).—Pseudopyrenula tessella (Pers.) Graff, Mycologia 9: 16 (1917); type: (L—holotype, lost).

Verrucaria ochroleuca Eschw., in Martius, Icon. Sel. Pl. Crypt. 2: 16, t. 8, figs 3, 4 (1828).—Pseudopyrenula ochroleuca (Eschw.) Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 209 (1890).—Spermatodium ochroleucum (Eschw.) Trevis., Conspect. Verruc.: 11 (1860).—Trypethelium ochroleucum (Eschw.) Nyl., Flora 52: 126 (1869); type: Brazil, Bahia, Caetité, Martius s. n. (M!—lectotype, Harris, Acta Amazon. (Suppl.) 14: 75 (‘1984’) [1986]).

Trypethelium duplex Fée, Ann. Sci. Nat., Bot. 23: 437 (1831).—Chooicia feei Trevis. (1861) nom. illeg.—Trypethelium cascarillae Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 30(3): 14 (1888) nom. illeg.—Pseudopyrenula duplex (Fée) Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 208 (1890); Bathelium duplex (Fée) C. W. Dodge, Ann. Missouri Bot. Gard. 40: 290 (1953); type: South America, “corticem Croton cascarillae” (G!—lectotype; Harris, Lichenogr. Thomsoniana: 142, 1998; BM!—isolectotype).

Trypethelium quassiicola Fée, Ann. Sci. Nat., Bot., sér. 1 23: 448 (1831); type: Jamaica, “in cortice Quassiae excelsae” (G!—lectotype, designated here; L—2 isolectotypes).

Trypethelium pallescens Fée, Ann. Sci. Nat., Bot., sér. 1 23: 440 (1831).—Trypethelium ochroleucum var. pallescens (Fée) Müll. Arg., Bot. Jahrb. Syst. 6: 392 (1885).—Pseudopyrenula ochroleuca var. pallescens (Fée) Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 209 (1890); type: Surinam, Weigelt s. n. (G!—lectotype, Müll. Arg., Bot. Jahrb. Syst. 6: 392, 1885; L!—isolectotype).

Trypethelium leprieurii Mont., Ann. Sci. Nat., Bot., sér. 2 19: 70 (1843); type: French Guiana, Leprieur s. n. (L—isotype).

Trypethelium triplex Nyl., Flora 52: 125 (1869); type: Brazil, Rio de Janeiro, Glaziou s. n. (H-Nyl—holotype, not seen).

Trypethelium euporum Kremp., Flora 59: 527 (1876); type: Brazil, Rio de Janeiro, Glaziou 6304 (M!—holotype).

Trypethelium bicolor var. pyrenuloides C. Knight, Trans. Proc. New Zealand Instit. 16: 405 (1884).—Trypethelium phlyctaena var. pyrenuloides (C. Knight) Zahlbr., Catal. Lich. Univ. 1: 498 (1922); type: New Zealand, Knight (WELT—holotype, not seen).

Trypethelium ochroleucum var. depauperatum Müll. Arg., Bot. Jahrb. Syst. 6: 392 (1885); type: Cuba, Müller, Verr. Cub. 571 (G!—lectotype, designated here; US—isolectotype, not seen).

Trypethelium leprosum Müll. Arg., Bot. Jahrb. Syst. 6: 393 (1885); type: Cuba, Wright s. n. (G!—lectotype, designated here; Müller, Verr. Cub. 155 p.p.).

Trypethelium subalbens Nyl., Flora 69: 178 (1886).—Bathelium subalbens (Nyl.) C. W. Dodge, Ann. Missouri Bot. Gard. 40: 290 (1953); type: São Tomé, Henriques s. n. (G!—lectotype, designated here; BM—isolectotype).

(Fig. 13F–L)

Thallus corticate, light greenish to brownish grey or yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic; pseudostromata 0·8–1·5(–2·0) mm broad., erumpent to prominent, covered by thallus but usually paler and cream-coloured. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 15–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone).

Distribution. Pantropical; reported from, for example, USA, Mexico, Puerto Rico, Cuba, Netherlands Antilles, Dominican Republic, Jamaica, Costa Rica, Nicaragua, Guyana, Surinam, French Guiana, Brazil, Uruguay, Ascension, São Tomé, Australia and New Zealand.

Discussion. In the complex traditionally labelled Trypethelium ochroleucum, the epithet phlyctaena is the oldest for specimens without hamathecium inspersion and with raised pseudostromata. This complex was even considered to include taxa with eccentric ostioles and hence was temporarily included in the synonymy of Astrothelium variolosum by Harris (1995), which is considered here a separate complex. Specimens previously labelled Trypethelium ochroleucum but with separate ascomata (not pseudostromatic) are assigned here to Astrothelium pulcherrimum.

New or confirmed country records. Nicaragua: Ometepe Island: Madera Volcan, 2001, Breuss 19160 (LI).—Netherlands Antilles: St. Eustatius: Quill, 1980, Sipman 15092 (B). Saba: Mt. Scenery, 1980, Sipman 15355 (B).—Puerto Rico: Ponce: Guanica, 1989, A. & M. Aptroot 25765 (ABL).—Dominican Republic: Monte Cristi: El Morro, 1987, Harris 19604 (ABL, NY).—Guyana: Upper Takutu: Dadanawa ranch, 1992, Sipman 57378 (B).—Uruguay: Rocha, Sierra de San Miguel, 1989, Osorio 9187 (hb. Kalb).—Ascension: Elliot’s Park, 1976, James s. n. (BM, ABL).

Astrothelium pleiostomum Redinger

Hedwigia 73: 58 (1933); type: Brazil, Amazonas, Santarem, Taperinha, Ginzberger (W—holotype, not seen).

Trypethelium violascens R. C. Harris, Acta Amazon. (Suppl.) 14: 78, fig. 11 (‘1984’) [1986]; type: Brazil, Amazonas, along Igarapé Caititu off Rio Uatumá, Buck 2937 (NY!—holotype; ABL!—isotype).

(Fig. 29I & J)

Thallus corticate, olive-green to yellowish, uneven somewhat bullate, often gall-like.

Ascomata pleurothelioid with eccentric, separate ostioles, diffusely pseudostromatic; pseudostromata 0·8–1·3 mm diam., erumpent, covered by thallus but with upper portion whitish, internally with yellowish pigment. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 28–33×8–10 µm, rather thin-walled, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−; pseudostroma internally K+ yellow, with anthraquinone.

Distribution. Neotropical (Brazil).

Discussion. This taxon is tentatively retained in Astrothelium, since the overall morphology and the ascospore type would also suggest placement in Viridothelium. The new synonymy of Trypethelium violascens is based on study of isotypes in ABL, which show that some characters are different from the description. The K-reaction is positive and the IKI-reaction is (at least now) absent; it was probably an artifact of fresh material.

Astrothelium porosum (Ach.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816721

Trypethelium porosum Ach., Synops. Lich: 106 (1814).—Verrucaria porosa (Ach.) Eschw., in Martius, Flora Brasil. 1: 135 (1833).—Leightonia porosa (Ach.) Trevis., Flora 44: 19 (1861).—Trypethelium sprengelii var. porosum (Ach.) Nyl., in Hue, Nouv. Archiv. du Muséum, sér. 3 4: 129 (1892).—Bathelium porosum (Ach.) C. W. Dodge, Ann. Missouri Bot. Gard. 40: 290 (1953); type: West Indies, “corticem Crotonis Cascarillae” (H-ACH 883!—holotype; S—isotype).

Trypethelium erubescens Kunze in Fée, Ann. Sci. Nat., Bot., sér. 1 23: 441 (1831).—Trypethelium ochroleucum var. erubescens (Kunze) Müll. Arg., Flora 68: 254 (1885); type: Surinam, Weigelt s. n. (L!—holotype).

Trypethelium brachysporum Malme, Ark. Bot. 19(1): 30 (1924).—Polymeridium brachysporum (Malme) Aptroot, in Aptroot & Cáceres, Nova Hedwigia 98: 11 (2014); type: Brazil, Mato Grosso, Cuyabá, Malme 2055 (S!—holotype; BM!—isotype).

(Fig. 12K & L)

Thallus corticate, light greenish to brownish grey or yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic; pseudostromata 0·8–1·5 mm diam., erumpent to promiment, covered by thallus but usually paler and cream-coloured. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 17–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone).

Distribution. Pantropical (see below).

Discussion. This taxon was until recently not distinguished from Astrothelium phlyctaena and A. pulcherrimum, and therefore its distribution is not well known. Together, these species have for a long time been labelled Trypethelium ochroleucum and under that name have been reported from the USA, Mexico, Bahamas, Jamaica, Cuba, Puerto Rico, Trinidad, Tobago, Costa Rica, El Salvador, Guyana, French Guiana, Surinam, Colombia, Venezuela, Galapagos, Brazil, Paraguay, Cape Verdes, Ascension, South Africa, Mozambique, Seychelles, India, Sri Lanka, China, Singapore, Philippines, Vietnam, Indonesia, Guam, Hawai’i, Northern Mariana Islands, New Caledonia, Papua New Guinea, Australia and New Zealand. Not all specimens on which these reports are based have been studied recently. Astrothelium porosum is the most common species of the three.

New country records. USA: Florida: Sanford, 1939, Johnson 3310 (ABL).—Mexico: Chiapas: Ocozocoautla, 1996, Wolf & Sipman s. n. (B). Guyana: Upper Takutu: Dadanawa ranch, 1992, Sipman 57358 (B).—French Guiana: Cayenne, 1993, Sipman 31532 (Lichenotheca Latinoamericana 149, ABL).—India: West Bengal: Sundarbans, Jhila, 2003, Jagadeesh Ram 13640 (ABL).—Indonesia: Irian Jaya: 11km W of Bupul, 1988, Hensen s. n. (ABL).—Papua New Guinea: Madang: Karkar, 1987, Aptroot 17623 (ABL).—Australia: Queensland: Cape Tribulation, 1983, Hale 64181 (ABL, US).

Astrothelium praetervisum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816722

Heufleria praetervisa Müll. Arg., Flora 68: 250 (1885).—Cryptothelium praetervisum (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 522 (1922); type: French Guiana, Leprieur 13 (G!—holotype).

(Fig. 37G)

Thallus corticate, brownish to often partly orange-brown and with orange pigment, mottled with whitish areas producing pycnidia, smooth to uneven.

Ascomata pleurothelioid, with eccentric, separate ostioles, pseudostromatic; pseudostromata 0·4–0·6 mm diam., immersed-erumpent, often forming lines, covered by thallus and partly with orange pigment except for blackish ostioles surrounded by whitish rim. Hamathecium clear. Ascospores 8 per ascus, muriform, fusiform, 33–68×12–22 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K+ partly purple, with anthraquinones.

Distribution. Neotropical (French Guiana).

Astrothelium pseudocyphellatum R. C. Harris

Acta Amazon. (Suppl.) 14: 62 (‘1984’) [1986]; type: Brazil, Amazonas, São Gabriel, Spruce 247 (BM!—isotype).

(Fig. 24F & G)

Thallus light greyish green, uneven to rugose, with superficial pockets of calcium oxalate crystals resembling pseudocyphellae.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 0·8–1·7 mm diam., prominent, covered by thallus usually paler or darker cream-coloured. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 23–27×7–8 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV+ yellow, K−, with lichexanthone.

Distribution. Neotropical (previously reported only from Brazil).

New country record. Guyana: Potaro-Siparuni: Kaieteur Falls National Park, 1996, Sipman 40458 (B).

Astrothelium pseudoplatystomum (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816723

Trypethelium pseudoplatystomum Makhija & Patw., Int. J. Mycol. Lichenol. 5: 244 (1992); type: Brazil, Heufler 130 (M—holotype, not seen).

Thallus corticate, green to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 1–2 mm diam., erumpent to prominent, with flattened top, covered by thallus except upper portion, with individual ascomata appearing dark and arranged in an astroid way. Hamathecium clear. Ascospores 8 per ascus, (5–)10–14-septate, fusiform, 29–52×8–12 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Brazil).

Astrothelium pseudovariatum (Upreti & Ajay Singh) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816724

Laurera pseudovariata Upreti & Ajay Singh, Bull. Jard. Bot. Nat. Belg. 57: 378 (1987); type: India, West Bengal, Howrah, Sibpur, Roychaudhury 675 (CAL—holotype, not seen).

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, diffusely pseudostromatic; pseudostromata 0·8–1·7 mm diam., prominent, hemispherical, covered by thallus. Hamathecium inspersed. Ascospores 8 per ascus, (sub-)muriform, 7–9×2–3-septate, 32–45×11–13 µm, fusiform, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (India).

Astrothelium puiggarii (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816725

Campylothelium puiggarii Müll. Arg., Flora 66: 245 (1883); type: Brazil, Rio de Janeiro, Puiggari 1460 (G!—holotype; NY!—isotype).

Campylothelium puiggarii var. pallescens Müll. Arg., Flora 66: 245 (1883); type: Brazil, Rio de Janeiro, Puiggari 1757 (G!—holotype).

(Fig. 38C & D)

Thallus corticate, olive-green, verrucose-bullate.

Ascomata pleurothelioid, with eccentric, separate ostioles, solitary, 0·8–1·3 mm diam., prominent to almost sessile, covered by thallus except for the protruding, papilliform, brownish grey ostiole, 1·0–1·5 mm diam. Hamathecium inspersed. Ascospores 2 per ascus, densely muriform, fusiform, 70–90×20–25 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Costa Rica and Brazil).

New country record. French Guiana: Saül, 1986, Montfoort & Ek 222 (B).

Astrothelium pulcherrimum (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816726

Trypethelium pulcherrimum Fée, Ann. Sci. Nat., Bot., sér. 1 23: 450 (1831).—Pseudopyrenula pulcherrima (Fée) Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 208 (1890); type: South America, “in America ad corticem Crotonis cascarillae” (G!—lectotype, designated here; L!—isolectotype).

Verrucaria diffluens Nyl., Ann. Sci. Nat., Bot., sér. 4 20: 252 (1863).—Pseudopyrenula diffluens (Nyl.) Müll. Arg., Flora 66: 248 (1883); type: Colombia, Bogotá, Lindig 2770 (H-Nyl 2211!—lectotype, designated here; BM!, BR—isolectotypes).

Trypethelium ochroleucum var. effusum Müll. Arg., Bot. Jahrb. Syst. 6: 392 (1885).—Pseudopyrenula ochroleuca var. effusa (Müll. Arg.) Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 209 (1890); type: Cuba, Wright s. n. (G!—lectotype, designated here; Müller, Lich. Cub., Ser. II: 566).

Trypethelium tricolor Müll. Arg., Bull. Soc. Roy. Bot. Belg. 32(1): 166 (1893); type: Costa Rica, Puntarenas, Boruca, Tonduz s. n. (G!—holotype; BR, US—isotypes; Pittier, Pl. Costaric. Exs. 6282).

Pseudopyrenula duplex var. simplicior Vain., Bol. Soc. Broteriana, Ser. 2 6: 177 (1930).—Trypethelium duplex f. simplicius (Vain.) Zahlbr., Catal. Lich. Univ. 10: 101 (1938).—Bathelium duplex f. simplicius (Vain.) C. W. Dodge, Ann. Missouri Bot. Gard. 40: 290 (1953); type: Mozambique, Palma, Pres de Lima 265 (TUR-Vain 34609—holotype, not seen).

Pseudopyrenula portoricensis J. Hedrick, Mycologia 22: 248 (1930); type: Puerto Rico, Mayagüez, Fink 1025 (MICH—holotype, not seen).

(Fig. 13B–D)

Thallus corticate, light olive-brown to brownish greenish grey to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary but usually dense, 0·2–0·3 mm diam., erumpent, covered by thallus except dark ostioles surrounded by whitish rim. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 15–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone).

Distribution. Pantropical, but distribution incompletely known because the species was not previously separated from Astrothelium phlyctaena and A. porosum (all taken together as Trypethelium ochroleucum s. lat.). Previously reported from Cuba, Puerto Rico, Colombia and Mozambique.

New country records. Guyana: Upper Takutu: Karanambo ranch, 1992, Sipman 57281 (B).—Indonesia: Java: Malang, 1939, Groenhart 5095 (ABL, L).

Astrothelium punctulatum Malme

Ark. Bot. 19(1): 14 (1924); type: Brazil, Matto Grosso, Santa Ana da Chapada, Malme, Lich. Regn. 2457 (S!—holotype; S!—isotype).

(Fig. 12G & H)

Thallus corticate, olive-green to greyish, verrucose-bullate, somewhat pseudogall-inducing.

Ascomata trypethelioid, with apical ostioles, solitary to confluent but often dense, 0·8–1·5 mm diam., immersed-erumpent in elevated thallus portions, covered by thallus except for dark ostiole surrounded by white ring. Hamathecium heavily inspersed. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 23–27×8–9 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ostioles UV+ yellow, K−, with lichexanthone.

Distribution. Neotropical (previously reported from Cuba and Brazil).

New country record. Guyana: Demerara-Berbice: Mabura Hill, 1988, Bleij & Biesmeijer s. n. (ABL). Upper Takutu: Kuyuwini Landing, 1992, Sipman 57021 (B).

Astrothelium pupula (Ach.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816727

Pyrenula pupula Ach., Synops. Meth. Lich.: 123 (1814).—Pseudopyrenula pupula (Ach.) Müll. Arg., Flora 68: 331 (1885).—Trypethelium pupula (Ach.) R. C. Harris, Lichenogr. Thomsoniana: 146 (1998); type: West Indies, s. col. (H-ACH 832!—holotype).

Pseudopyrenula porinoides Müll. Arg., Flora 69: 331 (1885).—Verrucaria porinoides Mont., Ann. Sci Nat., Bot. sér. 2 19: 59 (1843) non Ach.—Pseudopyrenula pupuloides M. Choisy, Mém. Soc. Bot. France 1953–1954: 58 (1954); type: French Guiana, Leprieur 731 (G!—lectotype, designated here; BM!—isolectotype).

(Fig. 14A & B)

Thallus corticate, greyish green to yellowish (lichexanthone colour), smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·4–0·6 mm diam., erumpent, covered by thallus except for blackish ostiolar area. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 33–40×10–13 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV+ yellow, K−. TLC: lichexanthone.

Distribution. Pantropical (previously reported from the West Indies, Costa Rica, French Guiana, Colombia, India and Sri Lanka).

New country records. Puerto Rico: Aguadilla: Bosque Estatal de Guajataca, 1989, Aptroot & Aptroot 25744 (ABL).—Guyana: East Demarara: Linden Highway, E of Timehri, 1985, Sipman & Aptroot 19587, 19602, 19558 (all ABL, B).—Venezuela: Bolivar: Cerro Guaiquinima, along Rio Carapo, 800 m, 1990, Sipman 27030 (B).—Ecuador: Morona-Santiago: Cordillera del Condor, 12km E of Los Encuentros, 1982, Aptroot 10442 (ABL).—Brazil: Minas Gerais: Serra do Caraça, Parque Natural do Caraça, 1250 m, 1997, Aptroot 40918 (ABL).—Malaysia: Sarawak: 1866, Beccari 188 p.p. (M, originally filed as syntype of Trypethelium leucostomum Kremp.).—Papua New Guinea: Central: Varirata National Park, 1987, Aptroot 19080 (ABL).

Astrothelium purpurascens (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816728

Heufleria purpurascens Müll. Arg., Bot. Jahrb. Syst. 6: 384 (1885).—Cryptothelium purpurascens (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 522 (1922); type: Cuba, Wright s. n. (G—holotype, not seen; NY!—isotype; Müller, Verr. Cub. 116b).

Cryptothelium rhodotitthon R. C. Harris, Acta Amazon. (Suppl.) 14: 65 (‘1984’) [1986]; type: Brazil, Roraima, along Manaus-Boa Vista Road, Buck s. n. (NY!—isotype).

(Fig. 37L)

Thallus corticate, olive-green to yellowish or reddish, uneven to bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 0·7–1·5 mm diam., immersed, completely covered by thallus except for protruding, dark red ostiole. Hamathecium clear. Ascospores 8 per ascus, densely muriform, fusiform, 100–130×25–30 µm, median septum thickened, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata with red, K+ green-reacting isohypocrellin.

Distribution. Neotropical (Brazil and Cuba).

Astrothelium pustulatum (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816729

Pseudopyrenula pustulata Vain., in Hiern, Cat. Afr. Pl. 2(2): 456 (1901).—Trypethelium pustulatum (Vain.) Zahlbr., Catal. Lich. Univ. 1: 499 (1922); type: Angola, Ambriz, Welwitsch 190 (TUR-Vain 30791!—holotype; BM!—isotype).

(Fig. 21G)

Thallus corticate, olive-green, uneven-verrucose, pseudogall-inducing.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic; pseudostromata 1–2 mm diam., prominent, completely covered by thallus, internally with yellow pigment, chiefly around the ostiolar region. Hamathecium inspersed. Ascospores 8 per ascus, 5–9-septate, 180–220×40–45 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances.

Distribution. African palaeotropical (Angola).

Discussion. In the original description a red pigment was mentioned, which is absent from the type material. The type of Trypethelium lageniferum Ach. also contains this taxon but additionally a species of Pyrenula, on which most of the description was based.

Astrothelium pyrenastrosulphureum Aptroot & Lücking nom. nov.

MycoBank No.: MB 816730

Pyrenastrum sulphureum Eschw., in Martius, Icon. Pl. Cryptog.: 17 (1828).—Astrothelium sulphureum (Eschw.) Müll. Arg., Flora 68: 255 (1885) nom. illeg. non Astrothelium sulphureum Nyl., Ann. Sci. Nat., Bot., sér. 4 20: 260 (1863); type: Brazil, Amazonas, Martius s. n. (M!—holotype).

Pyrenodium hypoxylon Fée, Essai Crypt. Écorc. Suppl.: 69 (1837).—? Astrothelium hypoxylon (Fée) Nyl., Mém. Soc. Sci. Nat. Cherbourg 5: 141 (1857); type not indicated.

Pyrenastrum sulphureum subsp. plicatum Eschw., in Martius, Flora Brasil. 1: 145 (1833); type: Brazil, Amazonas, Martius s. n. (M—holotype, not seen).

Astrothelium sulphureum var. subpallescens Nyl., Bull. Soc. Linn. Normand., sér. 2 2: 133 (1868); type: New Caledonia, Lifu, Deplanche s. n. (H-Nyl—holotype, not seen).

(Fig. 28A)

Thallus corticate, olive-green to yellowish, uneven to bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata irregularly pseudostromatic; pseudostromata 0·8–1·5 mm diam., erumpent to prominent, covered by thallus except for blackish ostiolar area. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 27–30×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances.

Distribution. Pantropical (Brazil, India and New Caledonia).

Astrothelium robustum Müll. Arg.

Bull. Soc. Roy. Bot. Belg. 32(1): 164 (1893); type: Costa Rica, Puntarenas, Rio Naranjo, 1893, Tonduz s. n. (G!—holotype; BR—isotype).

Astrothelium conigerum Zahlbr., Bull. Herb. Boissier, sér. 2 8: 459 (1908); type: Brazil, Amazonas, Rio Juruá, Ule 356 (W—holotype, not seen).

(Fig. 30H & I)

Thallus corticate, olive-green to yellowish, uneven to bullate, sometimes pseudogall-inducing.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to irregularly confluent or diffusely pseudostromatic, 1–3 mm diam., erumpent to prominent, covered by thallus except for blackish ostiolar area. Hamathecium clear. Ascospores 8 per ascus, (3–)5–7(–9)-septate, fusiform, (70–)90–130×20–30 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Brazil and Costa Rica).

New country records. Puerto Rico: Ponce: Caribbean National Forest, 1989, Aptroot & Aptroot 25603, 25602 (ABL).—Colombia: Nariño: Tumaco, Estacion Forestal La Espriella, 1986, Sipman & Velosa 33054 (B). Amazonas: Araracuara, opposite Isla Mariñame, 240 m, 1988, Sipman & Duivenvoorden 28328 (B).

Astrothelium rufescens (Müll. Arg.) Aptroot & Lücking comb. et stat. nov.

MycoBank No.: MB 816731

Trypethelium catervarium var. rufescens Müll. Arg., Bot. Jahrb. Syst. 6: 391 (1885); type: Cuba, Wright s. n. (G!—lectotype, designated here; FH-TUCK 3970!—isolectotype, designated here; Müller, Verr. Cub. 576).

(Fig. 18K & L)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·5–1·5 mm broad, immersed, forming irregular, whitish lines that become fused and reticulate. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 17–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (Cuba).

Distribution. This taxon belongs in the Astrothelium nitidiusculum aggregate, where it is characterized by its linear-reticulate, immersed, whitish pseudostromata.

Astrothelium sanguinarium (Malme) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816733

Laurera sanguinaria Malme, Ark. Bot. 19(1): 21 (1924).—Meristosporum sanguinarium Malme (1924) nom. inval; type: Brazil, Matto Grosso, Serra da Chapada da Buriti, Malme 2253 (S!—lectotype, designated here).

(Fig. 32G)

Thallus corticate, olive-green to yellowish, smooth to uneven to shallowly bullate.

Ascomata trypethelioid, with apical ostioles, distinctly pseudostromatic; pseudostromata 1–2 mm diam., erumpent to prominent, dark brown with pale brown ostioles, internally with bright to dark red pigment; pseudostromata up to 5 mm long and 2 mm broad. Hamathecium clear. Ascospores 8 per ascus, muriform, ellipsoid, (75–)90–130×15–25 µm, without strongly thickened median septum, hyaline, IKI+ violet.

Chemistry. Thallus UV−, K−; pseudostromata UV−, internally with red, K+ green-reacting isohypocrellin.

Distribution. Neotropical (Brazil).

Discussion. This taxon resembles a species of Bathelium but molecular data place it in Astrothelium. Specimens with whitish to pale cream-coloured pseudostromata represent an undescribed taxon labelled “Laurera Formosa” by R. C. Harris, the original material collected in Brazil by Malme (2514, S).

Astrothelium santessonii (Letr.-Gal.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816737

Laurera santessonii Letr.-Gal., Rev. Bryol. Lichénol. 37: 68 (1958); type: Ivory Coast, Mont Toukoui, cercle de Man, 14 August 1954, Santesson 10607d (UPS—holotype, not seen).

(Fig. 38B)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata pleurothelioid, with eccentric, separate ostioles, solitary to aggregate but not distinctly pseudostromatic, 0·7–1·5 mm diam., immersed-erumpent, covered by thallus except dark grey ostiolar area surrounded by irregular whitish rim. Hamathecium inspersed with red, K+ green droplets. Ascospores 8 per ascus, muriform, ellipsoid, 70–100×20–32 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−; hamathecium red, K+ green, with isohypocrellin.

Distribution. Pantropical (previously reported only from the Ivory Coast).

New country record. Guyana: Upper Takutu: Kuyuwini Landing, 1992, Sipman 58142 (B).

Astrothelium saxicola (Malme) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816738

Cryptothelium saxicola Malme, Svensk Bot. Tidskr. 30: 246 (1936); type: Brazil, Matto Grosso, Serra da Chapada, Bocca da Serra, Malme 2314 (S!—holotype).

(Fig. 39I)

Thallus pale greenish grey to yellowish, smooth to uneven or rough.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 1·0–1·8 mm diam., erumpent, applanately wart-shaped, completely covered by thallus, with upper part yellowish. Hamathecium clear. Ascospores 1 per ascus, densely muriform, ellipsoid, 215–230×60–80 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances.

Distribution. Neotropical (Brazil).

Discussion. This is one of the very few specimens of Trypetheliaceae encountered on rock, and one of the three species that seem restricted to rock; the others are Astrothelium stramineum, from the same locality, and Pseudopyrenula saxicola.

Astrothelium scoria (Fée) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816739

Trypethelium scoria Fée, Essai Crypt. Écorc.: 69 (1824).—Pseudopyrenula scoria (Fée) Vain., Bol. Soc. Broteriana, Ser. 2 6: 178 (1930); type: South America, “corticem Crotonis cascarillae” (G!—lectotype; Harris, Lichenogr. Thomsoniana: 147, 1998; G!, BM!—isolectotypes).

Pyrenula myriocarpa Fée, Essai Crypt. Écorc.: 74 (1824).—Verrucaria myriocarpa (Fée) Spreng., Syst. Veget. 4(1): 245 (1827).—Spermatodium myriocarpum (Fée) Trevis., Conspect. Verruc.: 11 (1860).—Trypethelium myriocarpum (Fée) Müll. Arg., Bot. Jahrb. Syst. 6: 391 (1885); type: South America, s. col. (G!—holotype).

Trypethelium scoria var. sordidius Nyl., Flora 52: 126 (1869); type: Brazil, Rio de Janeiro, Glaziou 1961 (H—holotype, not seen; M!—isotype).

Trypethelium dichroum Makhija & Patw., J. Hattori Bot. Lab. 73: 195 (1993); type: India, Andaman Islands, North Andaman, Diglipue Range, Millangram, 3 January 1986, Nagarkar & Patwardhan AMH 86.250 (ABL!—isotype).

Trypethelium indicum Makhija & Patw., J. Hattori Bot. Lab. 73: 199 (1993); type: India, Meghalaya, Willoe, 30 October 1977, Patwardhan & Nagarkar AMH 77.1130 (ABL!—isotype).

Trypethelium meghalayense Makhija & Patw., J. Hattori Bot. Lab. 73: 201 (1993); type: India, Meghalaya, near Sohararaim, on road to Cherapunji, 28 October 1977, Patwardhan & Nagarkar AMH 86.862 (ABL!—isotype).

(Fig. 18C–E)

Thallus corticate, olive-green to yellowish or brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic, 0·3–0·5 mm diam., erumpent, laterally covered by thallus but upper portion blackish brown with whitish rim. Hamathecium inspersed. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 15–27×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical (e.g. Puerto Rico, El Salvador, Guyana, French Guiana, Ecuador, Brazil, Zaire, India, China, Hong Kong, Papua New Guinea, Solomon Islands, New Caledonia).

Discussion. The epithet scoria is the oldest for the specimens with hamathecium inspersion of the complex previously known as Trypethelium nitidiusculum. As defined here, Astrothelium scoria is still heterogeneous and might comprise several species, but due to the limited material available we were unable to assess the variation of potential lineages represented by the synonyms listed here. The thallus of the newly synonymized Trypethelium indicum is an unspecific UV+ pinkish, not yellow as mentioned in the protologue.

New country records. El Salvador: Ahuachapán: Parque Nacional El Imposible, 1998, Sipman et al. 44570 (B).—Puerto Rico: Aguadilla: Guajataca, 1989, Aptroot & Aptroot 25737 (ABL).—Guyana: Upper Takutu: Dadanawa ranch, 1992, Sipman 57985 (B).—French Guyana: Regina: Camp Aratai, 2003, Sipman 50699 (B). Ecuador: Napo: Archidona, 1982, Aptroot & Hensen 10328 (ABL).—Zaire: Kivu: Equateur, Kalamba, 450 m, 1991, Müller s. n. (DR).—China: Yunnan: Xishuangbanna, 2003, Aptroot 57283 (ABL).—Hong Kong: Lantau: Ngong Ping, 2000, Aptroot 48638 (ABL).—Papua New Guinea: Madang: 11 km W of Brahman Mission, 1995, Sipman 38912 (B).—Solomon Islands: Santa Isabel: Tatamba, 1965, Hill 11170 (ABL, BM).—New Caledonia: Vieillard s. n. (BM).

Astrothelium scorioides Nyl.

Ann. Sci. Nat., Bot., sér. 5 7: 348 (1867); type: Colombia, Rio Negro, Lindig 132 (H-Nyl 120!—holotype; BM!—isotype).

(Fig. 27F & G)

Thallus corticate, pale greenish to yellowish, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to confluent, 0·3–0·5 mm diam., erumpent, conical with flattened top, covered by thallus and upper part with yellow-orange pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 33–45×10–17 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata with yellow to orange, K+ purple, UV+ red anthraquinone.

Distribution. Pantropical (previously reported from Venezuela, Colombia, Brazil and Seychelles).

New country records. Guyana: Upper Mazaruni: Mount Latipu, Sipman & Aptroot 19072 (ABL, B).—French Guiana: Saül, 1985, Aptroot 15442 (ABL, B).

Astrothelium scoriothelium Aptroot & Lücking nom. nov.

MycoBank No.: MB 816740

Trypethelium scorioides Leight., Trans. Linn. Soc. London 25: 459 (1866) non Astrothelium scorioides Nyl. (1867); type: Brazil, Amazonas, Spruce s. n. (BM!—holotype).

Pseudopyrenula infuscatula var. tecomae Vain., Ann. Acad. Sci Fenn., ser A 6(7): 197 (1915).—Trypethelium infuscatulum var. tecomae (Vain.) Zahlbr., Catal. Lich. Univ. 8: 128 (1931); type: Guadeloupe, Près du Camp-Jacob, Duss 1501 (TUR-Vain 30780!—holotype).

(Fig. 20C)

Thallus corticate, olive-green, uneven-verrucose.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata irregular, 0·7–1·7 mm broad, immersed to erumpent, exposed and blackish, with ostioles only visible by their pale rings. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 36–39×11–12 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances.

Distribution. Neotropical (reported from Guadeloupe, Venezuela and Colombia).

Astrothelium scorizum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816741

Trypethelium scorizum Müll. Arg., Bot Jahrb. Syst. 6: 389 (1885).—Trypethelium scorizum Nyl. (1876) nom. nud; type: Cuba, Wright s. n. (G!—holotype; BM!—isotype; Müller, Verr. Cub. 166a).

(Fig. 28C & D)

Thallus corticate, light olive-green to pale brownish, smooth to uneven-rugulose.

Ascomata pleurothelioid, with eccentric, separate ostioles, pseudostromatic; pseudostromata 0·7–1·5 mm diam., immersed-erumpent, forming irregular, in part confluent groups or lines, with the individual ascomata exposed and dark brown. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 20–26×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances.

Distribution. Neotropical (Cuba and Brazil).

New country record. Brazil: São Paulo: Pratânia, Fazenda Palmeira da Serra, 2009, Lücking 29814 (B, F).

Astrothelium sepultum Mont.

Ann. Sci. Nat., Bot., sér. 2 19: 74 (1843).—Cryptothelium sepultum (Mont.) A. Massal., Atti I. R. Istitut. Veneto, Ser. 3 5: 335 (1860).—Heufleria sepulta (Mont.) Trevis., Flora 44: 23 (1861).—Laurera sepulta (Mont.) O. E. Erikss., Op. Bot. 60: 50 (1981); type: French Guiana, Leprieur s. n. (BR, PC—syntypes).

Trypethelium connivens Nyl., Mém. Soc. Acad. Maine-et-Loire 4: 79 (1858).—Bathelium connivens (Nyl.) Trevis., Flora 44: 21 (1861).—Laurera connivens (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 503 (1922); type: Peru, s. col. (H-Nyl—holotype, not seen; PC!—isotype).

(Fig. 36H)

Thallus corticate, greyish green to pale yellowish brown (lichexanthone colour), smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to confluent but not distinctly pseudostromatic, 1·0–1·8 mm diam., erumpent, covered by thallus but with upper part whitish and ostiole dark. Hamathecium clear. Ascospores 2 per ascus, densely muriform, fusiform, 150–250×30–65 µm, with distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and usually also ascomata UV+ yellow, with lichexanthone.

Distribution. Neotropical (reported from Costa Rica, Guyana, French Guiana, Surinam, Brazil and Peru).

Astrothelium sierraleonense (C. W. Dodge) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816742

Tremotylium sierraleonense C. W. Dodge, Nova Hedwigia, Beih. 12: 104 (1964); type: Sierra Leone, Kori, Njala, Deighton M5693C (FH-DODGE!—holotype).

(Fig. 32C)

Thallus corticate, olive-green, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, dispersed to confluent but not distinctly pseudostromatic, 0·8–1·5 mm diam., erumpent, laterally covered by thallus but upper part conspicuously black and flattened, surrounded by an irregular whitish rim, internally with pockets of red crystals. Hamathecium clear. Ascospores 4 per ascus, muriform, ellipsoid, c. 80×14 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata externally UV−, K−; ascomata internally with red, K+ purple anthraquinone crystals.

Distribution. African palaeotropical (Sierra Leone).

Astrothelium sikkimense (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816743

Laurera sikkimensis Makhija & Patw., Mycotaxon 31: 580 (1988); type: India, Sikkim, Gangtok, Near Tangshi View Point, 17 September 1977, Patwardhan & Nagarkar AMH 77.1970 (ABL!—isotype).

(Fig. 33L)

Thallus corticate, brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 1–2 mm diam., immersed to erumpent, covered by thallus except black ostiole surrounded by whitish rim. Hamathecium inspersed. Ascospores 6–8 per ascus, muriform, ellipsoid, 120–140×25–30 µm, with somewhat thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (India).

Astrothelium spectabile (Aptroot & Ferraro) Aptroot & Lücking comb. nov.

MycoBank No.: MB 817086

Trypethelium spectabile Aptroot & Ferraro, Kurtziana 31: 59 (2005); type: Argentina, Misiones, Iguazú, 2003, Ferraro & Popoff 6602 (CTES!—holotype; ABL!—isotype).

(Fig. 21L)

Thallus corticate, light green, smooth.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·7–1·2 mm diam., immersed, completely covered by thallus. Hamathecium clear. Ascospores 8 per ascus, 11–19-septate, fusiform, 140–190×20–25 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (reported from Argentina).

Astrothelium sphaerioides (Mont.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816744

Trypethelium sphaerioides Mont., Ann. Sci. Nat., Bot., sér. 2 19: 73 (1843).—Bathelium sphaerioides (Mont.) Trevis., Spighe e Paglie: 20 (1861).—Laurera sphaerioides (Mont.) Zahlbr., Catal. Lich. Univ. 1: 506 (1922); type: French Guiana, Leprieur 80 (PC!—lectotype, designated here; H-Nyl!—isolectotype).

(Fig. 35H)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid to pleurothelioid, with apical to eccentric, separate ostioles, distinctly pseudostromatic; pseudostromata irregular, 0·5–1·5 mm broad, erumpent to prominent, whitish and black ostioles. Hamathecium clear. Ascospores 2–4 per ascus, muriform, ellipsoid, 80–175×25–50 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical; previously reported from Colombia (Isla de Providencia), French Guiana, Brazil and India.

New country records. El Salvador: Ahuachapán: Parque Nacional El Imposible, 1998, Welz s. n. (B).—Costa Rica: Guanacaste: 5 km NNW of Tilarán, 650 m, 2004, Aptroot 60659 (ABL, INB).—Panama: Veraguas: Bahía Honda, 2001, Cabrera & Etayo 18583 (ABL, hb. Etayo).—Puerto Rico: Mayagüez: Reserva Forestal Maricao, 800 m, 1989, Aptroot & Aptroot 24948, 24943 (ABL).—Guyana: Upper Takutu: Rupununi Savannah, Karanambo Ranch, 1992, Sipman 57321 (B).—Ecuador: Zamora-Chinchipe: Estacion Cientifico San Francisco, 40 km S of Loja, 2004, Sipman 52367 (B).

Astrothelium stramineum (Malme) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816745

Thelenella straminea Malme, Ark. Bot. 22A(1): 7 (1928).—Clathroporina straminea (Malme) Zahlbr., Catal. Lich. Univ. 8: 111 (1931).—Laurera straminea (Malme) P. M. McCarthy, Lichenologist 27: 348 (1995); type: Brazil, Matto Grosso, Serra da Chapada, Bocca da Serra, 15 March 1884, Malme 2550 (S!—holotype).

(Fig. 31D)

Thallus saxicolous, corticate, yellowish white, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary, 0·4–0·6 mm diam., hemispherical, covered by thallus. Hamathecium inspersed. Ascospores (2–)4(–6) per ascus, densely muriform, fusiform, 40–80×15–24 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone; ascomata UV−, K−.

Distribution. Neotropical (Brazil).

Discussion. This is one of the very few species of Trypetheliaceae ever encountered on rock, and one of three species that seem restricted to rock; the others are A. saxicola, from the same locality, and Pseudopyrenula saxicola.

Astrothelium straminicolor (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816746

Trypethelium straminicolor Nyl., Lich. Japon: 115 (1890); type: Malaysia, Labuan, Almquist s. n. (H-Nyl 174!—lectotype, designated here; S!—isolectotype).

Trypethelium stramineum Kremp., Nuov. Giorn. Bot. Ital. 7: 56 (1875), non Astrothelium stramineum (Malme) Aptroot & Lücking, see above; type: Sarawak, Beccari 212 (M!—lectotype, Makhija & Patwardhan, Int. J. Mycol. Lichenol. 5: 246, 1992).

Astrothelium subvariolosum Makhija & Patw., Biovigyanam 15: 64 (1989); type: India, Nicobar Islands, Great Nicobar, Campbell Bay to Laful Bay, 2 January 1987, Patwardhan & Nagarkar AMH 87-70 (ABL!—isotype).

(Fig. 27J–L)

Thallus corticate, olive-green to yellowish brown, uneven to shallowly bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 1–2 mm diam., erumpent to prominent, covered by thallus except for blackish upper part surrounded by a thin, whitish rim, with blackish ostiolar areas often arranged in lobate fashion. Hamathecium inspersed. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 20–25×7–9 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (India and Malaysia, including Sarawak).

Astrothelium subaequans Müll. Arg.

Bot. Jahrb. Syst. 6: 383 (1885); Astrothelium subaequans Nyl. (1876) nom. nud.; type: Cuba, Wright s. n. (G!—holotype; H-Nyl 108!—isotype; Müller, Verr. Cub. 146).

(Fig. 37H)

Thallus corticate, olive-green to yellowish, uneven to shallowly bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed, 1–2 mm diam., erumpent, covered by thallus except dark, protruding and papilliform ostiole, thinly covered with yellow-orange pigment. Hamathecium clear. Ascospores 8 per ascus, submuriform, fusiform, (45–)72–83×15–25 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; ascomata with yellow-orange, K+ purple, UV+ red anthraquinone.

Distribution. Neotropical (previously reported from Costa Rica and Cuba).

New country records. Venezuela: Bolivar: Cerro Guaiquinima, 1200 m, 1990, Sipman 26848 (B).—Guyana: Upper Mazaruni: N of Paruima Mission, Aymatoi savannah, 1000 m, 1997, Sipman 39806 (B). Potaro-Siparuni: Kaieteur Falls, around airstrip, 400 m, 1996, Sipman 40408 (B).

Astrothelium subcatervarium (Malme) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816747

Trypethelium subcatervarium Malme, Ark. Bot. 19(1): 29 (1924); type: Brazil, Matto Grosso, Serra da Chapada, Buriti, Malme s. n. (S!—holotype).

(Fig. 15L)

Thallus corticate, olive-green to yellowish brown, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to confluent or diffusely pseudostromatic, 0·4–0·6 mm diam., erumpent, whitish and dark brown ostiolar area thinly covered with yellow-orange pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 27–33×10–12 µm, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV+ orange, K+ red, with anthraquinone.

Distribution. Neotropical (reported from Cuba and Brazil). Incorrectly reported from Costa Rica: the specimen (Breuss 21057 in LI) proved to have K− pseudostromata.

Astrothelium subclandestinum Leight.

Trans. Linn. Soc. London 25: 460 (1866); type: Brazil, Amazonas, São Gabriel, Spruce 242 (BM!—holotype).

(Fig. 29L)

Thallus corticate, olive-green to yellowish, uneven to shallowly bullate.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata diffusely pseudostromatic; pseudostromata 1·0–1·5 mm diam., erumpent, laterally covered by thallus but apically greyish. Hamathecium inspersed. Ascospores 8 per ascus, 5-septate, fusiform, 50–75×14–22 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical (previously reported from Costa Rica, Brazil and Australia).

New country records. Venezuela: Bolivar: National Park Canaima, 2003, Berger 18510 (ABL, hb. Berger).—Guyana: Demerara-Berbice: Mabura Hill, 1985, Cornelissen 486 (ABL).—Papua New Guinea: Madang: Foothills of Finisterre Range, 1992, Aptroot 33209 (ABL).

Astrothelium subdiscretum (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816748

Trypethelium subdiscretum Nyl., Flora 52: 73 (1869).—Bathelium subdiscretum (Nyl.) Müll. Arg., Bot. Jahrb. Syst. 6: 395 (1885).—Laurera subdiscreta (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 506 (1922); type: India, West Bengal, Kurz s. n. (H-Nyl 322!—holotype; M! —isotype).

Bathelium phaeomelodes Müll. Arg., Bot. Jahrb. Syst. 6: 394 (1885).—Trypethelium phaeomelodes Nyl. (1876) nom. nud.— Laurera phaeomelodes (Müll. Arg.) Zahlbr., in Engler & Prantl, Natürl. Pflanzenfamil. 1(1*): 71 (1903); type: Cuba, Wright s. n. (G!—holotype; BM!—isotype; Müller, Verr. Cub. 170).

Laurera nigeriensis C. W. Dodge, Ann. Missouri Bot. Gard. 40: 297 (1953); type: Nigeria, Abadan, 1951, Thorold 165 (FH-Dodge—holotype, not seen).

Laurera subphaeomelodes Upreti & Ajay Singh, Bull. Jard. Bot. Nat. Belg. 57: 382 (1987); type: India, West Bengal, Howrah, Sibpur, Roychaudhury 649 (CAL—holotype, not seen).

Laurera verrucoaggregata Makhija & Patw., Mycotaxon 31: 586 (1988); type: India, Andaman Islands, North Andaman, Lamia Bay, 5 January 1986, Nagarkar & Sethy AMH 86.550 (ABL—isotype).

Laurera fusispora Makhija & Patw., Mycotaxon 31: 576 (1988); type: India, Karnataka, Anamod ghat, Anamod-Goa Road, 10 December 1974, Patwardhan & Prabhu AMH 74.2499 (ABL—isotype).

(Fig. 33A–E)

Thallus corticate, olive-green to brownish, uneven to verrucose-rugulose.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·8–1·5(–2·0) mm broad, immersed-erumpent, irregular to linear and often confluent and reticulate, with ascomata exposed and black. Hamathecium inspersed, clear. Ascospores 8 per ascus, small muriform, ellipsoid, 35–50×12–18 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Pantropical (previously reported from Cuba, Costa Rica, Nigeria, India and Thailand).

New country records. Netherlands Antilles: Bonaire: Ceru Kammia, 1977, van Slageren 8406-b (ABL).—Dominican Republic: La Altagracia, Bayahibe, 1981, Buck 5141 (NY, hb. Kalb).—Colombia: Caqueta, Araracuara, opposite airstrip, 250 m, 1988, Sipman & Duivenvoorden 27909 (B).—Venezuela: Bolivar: Cerro Guaiquinima, along Rio Carapo, 800 m, 1990, Sipman 26942 (B).—Guyana: between Karanambo and Kwaimatta, 1988, Maas et al. 7709d (ABL).—Sri Lanka: Anuradhapura: Wipattu, Santesson 26185 (S).—Singapore: Pulau Ubin Island: 2000, Sipman & Tan 46340 (B).

Astrothelium subdisjunctum (Müll. Arg.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816749

Bottaria subdisjuncta Müll. Arg., Bot. Jahrb. Syst. 6: 395 (1885).—Laurera subdisjuncta (Müll. Arg.) R. C. Harris, Acta Amazon. (Suppl.) 14: 66 (‘1984’) [1986]; type: Cuba, Wright s. n. (G!—holotype; Müller, Verr. Cub. II: 676).

Laurera dodgei Letr.-Gal., Rev. Bryol. Lichénol. 27: 71 (1958); type: Costa Rica, Puntarenas, Rio Sandalo, Galjo Dulce, Peninsula de Osa, 0–10 m, 23 August 1936, Dodge & Goerger 10063 (FH-DODGE!—holotype).

Clathroporina wainiana Zahlbr., Sitzungsber. K. Akad. Wiss. Wien, math.-naturw. Classe 111(1): 364 (1902); type: Brazil, Rio de Janeiro, Höhnel 151 (W—holotype, not seen).

(Fig. 35I & J)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata 0·7–1·5 mm broad, immersed-erumpent, irregular to linear and often confluent and reticulate, with ascomata exposed and black. Hamathecium clear. Ascospores 4 per ascus, densely muriform, fusiform, 120–175×30–45 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from the USA, Cuba, Costa Rica, Guyana, Venezuela and Brazil).

New country records. French Guiana: Cayenne: Cayenne, along forest track called Risque tout, 50 m, 1985, Aptroot 15126 (ABL, B). Saint-Laurent-du-Maroni: Saül, 1985, Aptroot 15440 (ABL, B); Saül, 1986, Montfoort & Ek 290 (ABL).

Astrothelium subdissocians (Nyl. ex Vain.) Aptroot & Lücking comb. et stat. nov.

MycoBank No.: MB 816750

Pseudopyrenula ochroleuca var. subdissocians Nyl. ex Vain., Bot. Tidskr. 29: 147 (1909).—Trypethelium ochroleucum var. subdissocians (Vain.) Hue (1892) nom. nud.—Trypethelium pallescens var. subdissocians Nyl. (1864) nom. nud; type: Thailand, Koh Chang, Schmidt 11 (TUR-Vain 30761!—holotype).

(Fig. 28L)

Thallus corticate, light olive-green to yellowish brown, smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed to irregularly confluent, 0·4–0·6 mm diam., erumpent to prominent, with complete whitish cover. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 20–25×7–9 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Palaeotropical (reported from Thailand).

Astrothelium subfuscum Kremp.

Nuovo Giorn. Bot. Ital. 7: 64 (1875); type: Singapore, Beccari 236 (M!—holotype).

(Fig. 28H & I)

Thallus corticate, olive-green to brownish, rather thick, uneven to bullate-folded.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata confluent to diffusely pseudostromatic; pseudostromata 0·7–1·3 mm diam., erumpent, irregular in outline, laterally covered by thallus but upper part of ascomata exposed, blackish and groups of ascomata bordered by thin, whitish line. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 21–27×8–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (previously reported from India, Singapore, Sabah, New Caledonia and Papua New Guinea). Reports from the Neotropics refer probably to Astrothelium crassum.

New country records. China: Hong Kong: New Territories, Tai Po Kau Nature Reserve, 1998, Aptroot 43290 (ABL).—Philippines: Luzon: Sorsogon, Irosin, 1916, Elmer 17111 (B).

Astrothelium superbum (Fr.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816751

Trypethelium superbum Fr., Syst. Orb. Veget. 1: 287 (1825) .— Campylothelium superbum (Fr.) Müll. Arg., Flora 68: 251 (1885); type: Indonesia, s. col. (UPS—holotype, not seen).

Thallus corticate, olive-green, smooth to uneven.

Ascomata pleurothelioid, with eccentric, separate ostioles, solitary to irregularly grouped, 1–2 mm diam., immersed-erumpent, covered by thallus. Hamathecium clear. Ascospores 8 per ascus, densely muriform, fusiform, 130–165×35–45 µm, without distinctly thickened median septum, hyaline or becoming yellowish, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (reported from India and Indonesia).

Astrothelium tenue (Aptroot) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816752

Campylothelium tenue Aptroot in Aptroot et al., Biblioth. Lichenol. 98: 53. (2008); type: Costa Rica, Alajuela, Volcán Tenorio National Park, surroundings of Pilón Biological Station, Sipman 51867 (B!—holotype).

(Fig. 38K & L)

Thallus corticate, olive-green, smooth to uneven.

Ascomata pleurothelioid, with eccentric, separate ostioles, 0·4–0·7 mm diam., erumpent, completely covered by thallus. Hamathecium clear. Ascospores 4–8 per ascus, muriform, fusiform, 95–115×20–28 µm, with distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Possibly pantropical (so far reported only from Costa Rica).

Discussion. This species is characterized by the fusiform distoseptate ascospores, in which the stellate primary locules remain recognizable in the mature ascospores.

New country records. Guyana: Demerara-Berbice: Mabura Hill, 1988, Bleij & Biesmeijer (ABL).—Singapore: Nee Soon, 2000, Sipman et al. 46164 & 46159 (B).

Astrothelium thelotremoides (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816753

Verrucaria thelotremoides Nyl., Ann. Sci. Nat., Bot., sér. 5 7: 346 (1867).—Pseudopyrenula thelotremoides (Nyl.) Müll. Arg., Flora 66: 248 (1883).—Trypethelium thelotremoides (Nyl.) R. C. Harris., Lichenogr. Thomsoniana: 148 (1986); type: Colombia, Rio Negro, Lindig 51 (H-Nyl 1113!—lectotype, designated here; BM!—isolectotype).

(Fig. 14D & E)

Thallus corticate, olive-green to brownish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, solitary to irregularly confluent, 0·5–0·8 mm diam., erumpent, covered by thallus with ostiolar area dark greyish with thin whitish rim, internally with red pigment. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 50–65×15–21 µm, hyaline, IKI−.

Chemistry. Thallus and ascomata UV−, K−; ascomata internally K+ purple, with an anthraquinone.

Distribution. Neotropical (previously reported only from Colombia).

Discussion. The type sheet contains three specimens; two are Ocellularia species and the third specimen is designated here as lectotype.

New country record. Brazil: Bahia: Entre Rios, 1981, Boom & Mori 1010 (hb. Kalb, NY).

Astrothelium trypethelizans (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816754

Verrucaria trypethelizans Nyl., in Crombie, J. Linn.Soc. London, Bot. 20: 60 (1883).—Pseudopyrenula trypethelizans (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 359 (1922).—Trypethelium trypethelizans (Nyl.) R. C. Harris, Lichenogr. Thomsoniana: 148 (1998); type: Malaysia, Malacca, Tanjong, Maingay 149 (BM!—lectotype; Harris, Lichenogr. Thomsoniana: 148, 1998).

(Fig. 21D)

Thallus corticate, olive-green to yellowish brown, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, dispersed to largely confluent, 0·3–0·5 mm diam., erumpent with flattened top, brownish black. Hamathecium clear. Ascospores 8 per ascus, 7–9-septate, fusiform, 32–38×10–11 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Eastern palaeotropical (reported from Malaysia).

Astrothelium tuberculosum (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816755

Pseudopyrenula annularis var. tuberculosa Vain., Ann. Acad. Sci Fenn, ser. A 6(7): 196 (1915).—Trypethelium crassum var. tuberculosum (Vain.) Zahlbr., Catal. Lich. Univ. 8: 128 (1932).—Trypethelium tuberculosum (Vain.) R. C. Harris, Acta Amazon. (Suppl.) 14: 76 (‘1984’) [1986]; type: Guadeloupe, Safraga, Duss 1415 (TUR-Vain 30778!—holotype).

(Fig. 20L)

Thallus corticate, olive-green to brownish, strongly bullate.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic; pseudostromata 1–2 mm diam., prominent to sessile, irregular, fully covered by thallus except dark ostioles surrounded by whitish rim. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform, 48–65×17–20 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Guadeloupe, Trinidad, Tobago, Costa Rica, Guyana, French Guiana, Venezuela, Galapagos and Colombia).

New country record. Brazil: São Paulo: Campos de Jordão, 1800 m, 1997, Sipman 41127 (SP, B).

Astrothelium ubianense (Vain.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816756

Pseudopyrenula ubianensis Vain., Ann. Acad. Sci. Fenn., ser. A 15(6): 353 (1921).—Trypethelium ubianense (Vain.) Zahlbr., Catal. Lich. Univ. 8: 129 (1932); type: Philippines, Ubian, Merrill 5403 p.p. (TUR-Vain 30798!—holotype).

(Fig. 21C)

Thallus corticate, light olive-green to yellowish, smooth to uneven.

Ascomata trypethelioid, with apical ostioles, pseudostromatic; pseudostromata irregular, 0·8–1·5 mm broad, basally covered by thallus but otherwise whitish, with the ostiolar areas of individual ascomata black and strongly contrasting. Hamathecium clear. Ascospores 8 per ascus, 5–11-septate, fusiform, 30–36(–49)×7–10 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances reported.

Distribution. Eastern palaeotropical (reported from India and the Philippines).

Astrothelium variatum (Nyl.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816757

Trypethelium variatum Nyl., Ann. Sci. Nat., Bot., sér. 5 7: 347 (1867).—Laurera variata (Nyl.) Zahlbr., Catal. Lich. Univ. 1: 507 (1922); type: Colombia, Rio Negro, Lindig s. n. (H-Nyl!—holotype).

Heufleria subvariata Müll. Arg., Bot. Jahrb. Syst. 6: 384 (1885).—Trypethelium subvariatum Nyl. (1876) nom. nud.— Cryptothelium subvariatum (Müll. Arg.) Zahlbr., Catal. Lich. Univ. 1: 523 (1922); type: Cuba, Wright s. n. (G!—holotype; Müller, Verr. Cub. 164).

(Fig. 35A & B)

Thallus corticate, olive-green, smooth to uneven or shallowly rugose.

Ascomata trypethelioid, with apical ostioles, diffusely pseudostromatic; pseudostromata 0·7–1·4 mm diam., prominent, irregular, covered by thallus except dark ostioles. Hamathecium clear. Ascospores 8 per ascus, small muriform, fusiform, 24–35×11–13 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV−, K−. TLC: no substances detected.

Distribution. Neotropical (previously reported from Cuba and Colombia).

New country record. Venezuela: Amazonas: Alto Orinoco, 15 km W of Esmeralda, 110 m, 1997, Hafellner & Komposch 209-3-7 (GZU).

Astrothelium variolosum (Ach.) Müll. Arg.

Flora 68: 255 (1885).—Trypethelium variolosum Ach., Synops. Lich.: 104 (1814); type: “ad corticem Chinchonae flavae” (H-ACH 877!—holotype; S!—isotype).

Trypethelium papillosum Ach., Synops. Lich.: 104 (1814) non Knight (1886).—Bathelium papillosum (Ach.) C. W. Dodge, Ann. Missouri Bot. Gard. 40: 288 (1953); type: Guinea, s. col. (H-ACH 878A!—holotype; S!—isotype).

Pyrenula epapillata Fée, Essai Crypt. Écorc.: 78 (1824).—Spermatodium epapillatum (Fée) Trevis., Conspect. Verruc.: 11 (1860); type (PC—syntypes, not seen).

Trypethelium papillosum var. fuscum Müll. Arg., Flora 68: 255 (1885); type: French Guiana, Leprieur 467 p.p. (G!—holotype).

(Fig. 24A–D)

Thallus corticate, pale greenish grey to pale yellowish brown (lichexanthone colour), smooth to uneven.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata pseudostromatic; pseudostromata 0·7–1·5 mm broad, erumpent to prominent, irregular in outline, covered by thallus but often paper to whitish, with dark ostioles. Hamathecium clear. Ascospores 8 per ascus, 3-septate, fusiform-ellipsoid, 20–26×7–9 µm, hyaline, IKI−.

Chemistry. Thallus and pseudostromata UV+ yellow, K−, with lichexanthone (or 1,8-dihydroxy-3,6-dimethoxyxanthone).

Distribution. Pantropical (previously reported from the continental USA, Bahamas, Cuba, Trinidad, Costa Rica, French Guiana, Venezuela, Colombia, Brazil, Bolivia, Sri Lanka, Thailand, Guam, Northern Mariana Islands and Australia).

Discussion. Further synonyms in the genera Verrucaria and Spermatodium are mentioned by Zahlbruckner (1922), but not cited here as the species was used in a different sense at that time and the material has not been checked. Astrothelium variolosum is accepted here in a very much more restricted sense than by Harris (1995), who included specimens with or without lichexanthone, with apical or eccentric ostioles, with simple or aggregated ascomata and with or without hamathecium inspersion, each of which are species characters in our concept. Specimens which differ only in an inspersed hamathecium are known from, for example, Guyana (Sipman 58039, 39954, and 39969). A new species is not described for them because it cannot be ruled out that one of the taxa that have been synonymized with A. variolosum has an inspersed hamathecium.

New country records. Puerto Rico: Aguadilla: Bosque Estatal de Guajataca, 1989, Aptroot & Aptroot 25730 (ABL).—Guyana: Upper Demerara-Berbice: Mabura Hill, 1994, DePriest et al. 9086 (B).—Indonesia: Java: Malang, 1932, Groenhart 412 (ABL, L).—Papua New Guinea: Madang: Madang, 1992, Aptroot 30078 (ABL).

Astrothelium versicolor Müll. Arg.

Flora 71: 495 (1888); type: Puerto Rico, Sintenis 6 (G!—holotype).

(Fig. 23C)

Thallus corticate, whitish to pale yellowish, strongly bullate-folded.

Ascomata astrothelioid, several chambers joined with eccentric, fused ostioles; joined ascomata dispersed but dense, 0·2–0·4 mm diam., completely immersed in thallus folds and warts, visible only by their darker ostioles thinly furnished with yellow-orange pigment. Hamathecium clear. Ascospores 8 per ascus, fusiform-ellipsoid, 3-septate, 28–35×10–15 µm, hyaline, IKI−.

Chemistry. Thallus UV+ yellow, K−, with lichexanthone; ascomata apically K+, UV+ red, with anthraquinone.

Distribution. Pantropical (previously reported from the USA, Bahamas, Cuba, Puerto Rico, Trinidad, Costa Rica, Guyana, French Guiana, Venezuela, Brazil, Bolivia, Philippines and New Caledonia).

New country record. Papua New Guinea: Central: Varirata National Park, 1995, Aptroot 39772 (ABL).

Astrothelium vezdae (Makhija & Patw.) Aptroot & Lücking comb. nov.

MycoBank No.: MB 816758

Laurera vezdana Makhija & Patw., Mycotaxon 31: 587 (1988) [as ‘vezdae`]; type: India, Maharashtra, Vishalgarh, Amba-Gajapur Road, 6 December 1974, Nagarkar & Prabhu AMH 74.2183 (ABL!—isotype).

(Fig. 32K)

Thallus corticate, olive-green to brownish, uneven-rugulose.

Ascomata trypethelioid to pleurothelioid, with apical to eccentric, separate ostioles, pseudostromatic; pseudostromata 1·0–1·3 mm diam., erumpent to prominent, covered by thallus but thinly covered with orange pigment. Hamathecium clear. Ascospores 2–4 per ascus, muriform, ellipsoid, 80–175×25–50 µm, without distinctly thickened median septum, hyaline, IKI−.

Chemistry. Thallus UV−, K−; pseudostromata UV+ reddish, K+ purple, with orange anthraquinone.

Distribution. Possibly pantropical (previously reported only from India).

New country record. Guyana: Upper Takutu: c. 35 km S of Aishalton, 250 m, 1992, Sipman 51722 (B).

Bathelium Ach.

Methodus: 111 (1803); type: Bathelium mastoideum Ach. (holotype).

Riddlea C. W. Dodge, Annals of the Missouri Botanical Garden 40: 287 (1953); type: Riddlea papillosa C. W. Dodge (= Bathelium cf. madreporiforme, holotype).

Thallus corticate, olive-gree