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Taxonomic studies on 

members of thelotrematoid Ostropales 

(lichenized Ascomycota) 

in Australia 

Inaugural-Dissertation 

zur 

Erlangung des Doktorgrades 

Dr. rer. nat. 

des Fachbereichs 

Biologie und Geografie 

an der 

Universität Duisburg-Essen 

vorgelegt von 

Armin Mangold 

aus Stuttgart 

Januar 2008

Die der vorliegenden Arbeit zugrunde liegenden Experimente wurden am 

Institut für Botanik der Universität Duisburg-Essen oder an einer anderen 

gleichwertigen Einrichtung durchgeführt. 

1. Gutachter: PD Dr. H. T. Lumbsch 

2. Gutachter:Prof. Dr. F. Kauff 

3. Gutachter: 

Vorsitzender des Prüfungsausschusses: Prof. Dr. Sures 

Tag der mündlichen Prüfung: 13.08.2008

ACKNOWLEDGEMENTS 3 

ACKNOWLEDGEMENTS 

I would like to acknowledge the help, support and interest of all friends, colleagues and institutions 

during the last five years. 

First of all, I wish to thank all the private collectors and herbaria for providing material for my project. 

These were the herbaria ABL, B, BCRU, BG, BM, BP, BRI, C, CANB, COLO, DUKE, E, F, FH, G, 

GLAM, GZU, H, HIRO, HO, L, M, MEL, MSU, NSW, NY, PERTH, RAMK, S, TNS, TUR, UPS, 

US, WELT, WIS and the private herbarium of Klaus Kalb (Neumarkt, Germany), who is also thanked 

for many helpful information. James C. Lendemer (Philadelphia) is not only thanked for providing 

fresh material for the molecular analyses but also for the many stimulating discussions. My particular 

appreciation for the support and hospitality during my visits go to the Wisconsin State Herbarium, 

University of Wisconsin (Madison), Paula DePriest and her co-workers at the U.S. National 

Herbarium, Smithsonian Institution (Washington DC) and, last but not least, to Ulrik Søchting and the 

National Museum of Natural History of Denmark, University of Copenhagen. I am also particularly 

indebted to Scott LaGreca, Lichen Curator at the Natural History Museum in London, for his 

invaluable help and patience when requesting an almost endless amount of type material. The visited 

National Parks of Queensland, New South Wales and Victoria and the administrative institutions of 

these states are hereby thanked for permitting the collecting. 

Barbara Emmerer (Graz), Andreas Frisch (Regensburg) and Gintaras Kantvilas (Hobart) are thanked 

for the helpful information and comments throughout the project. The Royal Botanical Garden Madrid 

and all associated workers, but especially María P. Martín, is cordially thanked for the hospitality and 

the great help and support during my stay in Madrid. The molecular analyses in their final extent 

would have been impossible without their encouragement. A special thanks goes to Jack Elix 

(Canberra), not only for his ceaseless support and help during this project, but also for his kindhearted 

hospitality and his company during our visit to Canberra, for his hints and suggestions regarding 

collecting sites and for enabling two fruitful field trips in New South Wales. 

I am also much obliged to the botany working group at the University Duisburg-Essen and for the very 

instructive time I was able to spent there. Particularly Stefanie Kautz is furthermore thanked for her 

help with the lab work and for her brilliant logistical superintendence of all transatlantic matters. 

Most of all, however, I am indebted to the Field Museum in Chicago and all the people involved. 

During the more than four years of my stay, this marvelous place has become a second home. Above 

all, I am grateful towards my supervisor Thorsten Lumbsch, who was not only the initiator of this 

project, but more so led to the migration into the New World. He was always a competent, reliable and 

patient supporter of my efforts to become a lichenologist, and I appreciate his friendship and, amongst 

others, the fact that he endured my driving abilities and my company in a tiny campervan in the 

Australian wilderness for almost four weeks. A special thanks goes to my fellow combatants and 

former roommates Imke Schmitt and Nora Wirtz for their excellent collegiality and friendship, but 

also for the linguistic corrections of the manuscript. All staff of the Botany Department is sincerely 

thanked, in particular Eimy Rivas Plata, who did a great job with a last minute photo session, and 

Mike Dillon, who provided additional funding for an extension of the project. 

Finally, I would like to thank my wonderful wife, co-fieldworker and image editor Anne Lass for her 

love, tolerance, support, encouragement and help over all this years. 

This study was financially supported by a grant of the Australian Biological Resources 

Studies (ABRS), the curator start-up fund of Thorsten Lumbsch, a fund of the Botany 

Department, the Women’s Board funding (the latter three all Field Museum, Chicago) and the 

SYNTHESYS program of the European Union (ES-TAF).

ABSTRACT 4 

ABSTRACT 

A taxonomic revision of the Australian members of trentepohlioid, ecolumellate 

thelotremataceaen Graphidaceae (Ostropales, lichenized Ascomycota) is presented. This 

species-rich group of crustose, predominantly tropical and subtropical lichens has been 

neglected for a long time, and is still poorly known. Particularly the studies for Australia are 

highly fragmentary. 

Approximately 4000 specimens, including a total of c. 500 type specimens, herbaria 

collections, and almost 1000 samples collected during two field trips, were examined 

morphologically, anatomically, and chemically. A total of 103 species of 13 genera (including 

one newly described genus) could be identified. 33 species are new records for Australia, and 

20 species were described new to science. Of the currently listed 71 names of this group for 

Australia, 50 are accepted as valid species. The remaining names and species, listed in an 

appendix, proved to belong to other genera, be synonymous to other taxa or falsely identified 

collections. 21 species (including species not known for Australia) are newly combined and 

several new synonyms are proposed. 

In the taxonomic part an introduction is provided, including a historical overview of the 

development of generic concepts and delimitations within the treated group. This is followed 

by a brief presentation of earlier taxonomic studies and collections of thelotrematoid lichens 

in Australia. It also provides a general treatment of the climatic and biogeographical 

conditions of the continent. Further, the main characters are described, including anatomical, 

morphological and chemical features. The ecology and distribution of thelotrematoid lichens 

are discussed. All treated species are described and illustrated, their distribution is mapped, 

and a key for the identification provided. An overview of the distinguishing characters 

between genera is presented in a table. 

To illuminate the phylogenetic relationships within this group, three phylogentic analyses 

were carried out using partial sequences of nuclear and mitochondrial ribosomal DNA loci. 

The first analysis employed a combined dataset of the mitochondrial SSU (mtSSU) and the 

nuclear LSU (nuLSU) of 105 species of Ascomycetes. As a result, it could be shown that the 

hitherto separted two families of Ostropales, Graphidaceae and Thelotremataceae, are not 

monophyletic, but form several lineages within one strongly supported monophyletic group. 

Consequently, Thelotremataceae is reduced to synonymy with Graphidaceae. The second and 

third analyses were carried out using a single dataset of nuLSU and mtSSU sequences 

respectively. In the nuLSU analysis 40 specimens of 38 species were included, in the mtSSU 

analysis 50 specimens of 25 species were examined. These analyses showed that all genera of 

thelotrematoid lichens included in the study are para- or polyphyletic, with the sole exception 

of Diploschistes, and that the present generic concept within Graphidaceae needs revision. 

The new genus Melanotopelia is introduced based on molecular and morphological evidence.

TABLE OF CONTENTS 5 

TABLE OF CONTENTS 

0. Deutschsprachige Zusammenfassung: Taxonomische Studien von Mitgliedern 

der thelotrematoiden Ostroplaes (lichenisierte Ascomycota) in Australien ................ 9 

1. Introduction ................................................................................................................. 13 

2. Taxonomic part ............................................................................................................ 15 

2. 1. Historical overview ............................................................................................... 15 

2. 2. Taxonomic studies and collections of trentepholioid thelotremataceaen 

Graphidaceae in Australia ..................................................................................... 16 

2. 3. Geography of Australia ......................................................................................... 17 

2. 4. Materials and Methods .......................................................................................... 19 

2. 4. 1. Examined collections ...............................................................................19 

2. 4. 2. Methods .................................................................................................. 20 

2. 4. 3. Citations .................................................................................................. 20 

2. 4. 4. Abbreviations .......................................................................................... 20 

2. 5. Main characters of trentepholioid thelotremataceaen Graphidaceae ....................... 21 

2. 5. 1. Thallus .................................................................................................... 21 

2. 5. 2. Ascomata ................................................................................................ 22 

2. 5. 3. Conidiomata ............................................................................................ 25 

2. 5. 4. Chemistry ................................................................................................ 26 

2. 6. Ecology ................................................................................................................ 28 

2. 6. 1. Substrate ................................................................................................. 28 

2. 6. 2. Habitat .................................................................................................... 29 

2. 7. Distribution ........................................................................................................... 29 

2. 8. Key to the treated species ...................................................................................... 32 

2. 9. Genera ................................................................................................................. 42 

2. 9. 1. Chapsa .................................................................................................... 44 

2. 9. 2. Chroodiscus ............................................................................................ 85 

2. 9. 3. Fibrillithecis ............................................................................................ 89 

2. 9. 4. Leptotrema .............................................................................................. 93 

2. 9. 5. Leucodecton ............................................................................................ 96 

2. 9. 6. Melanotopelia ....................................................................................... 111 

2. 9. 7. Myriotrema ........................................................................................... 114 

2. 9. 8. Nadvornikia .......................................................................................... 156 

2. 9. 9. Pseudoramonia ..................................................................................... 159 

2. 9. 10. Reimnitzia ............................................................................................. 161 

2. 9. 11. Thelotrema............................................................................................. 164 

2. 9. 12. Topeliopsis ............................................................................................ 255 

2. 9. 13. Species of uncertain taxonomic placement 

(Leptotrema schizoloma-group) ............................................................. 281 

3. Molecular phylogeny .................................................................................................. 294 

3. 1. Introduction ........................................................................................................ 294 

3. 2. Materials and methods ........................................................................................ 294 

3. 2. 1. Material ................................................................................................. 294 

3. 2. 2. DNA extraction ..................................................................................... 299 

3. 2. 3. PCR amplification and purification of PCR products ............................. 299


3. 2. 4. Sequencing and sequence assembly ....................................................... 299 

3. 2. 5. Sequence alignments ............................................................................. 300 

3. 2. 6. Phylogenetic analysis ............................................................................ 300 

3. 3. Results ............................................................................................................... 301 

3. 3. 1. Analysis 1 ............................................................................................. 301 

3. 3. 2. Analysis 2 ............................................................................................. 303 

3. 3. 3. Analysis 3 ............................................................................................. 305 

3. 4. Discussion .......................................................................................................... 307 

3. 4. 1. Phylogenetic relationship of Graphidaceae and Thelotremataceae 

(analysis 1) ............................................................................................ 307 

3. 4. 2. Phylogenetic relationship of genera within Graphidaceae 

(analysis 1, 2 and 3)................................................................................ 308 

4. References ............................................................................................................... 311 

5. Appendices ............................................................................................................... 322 

5. 1. Tentatively identified species............................................................................... 322 

5. 2. Excluded or dubious names ................................................................................. 322 

5. 3. List of synonyms ................................................................................................. 325 

Tables 

Table 1: TLC characteristics of the four hitherto unknown substances (according to Elix, pers. com.) 

and for the ‘cinchonarum und platysporum unknown’ ............................................................. 28 

Table 2: Australian distribution types of the treated species ................................................................... 31 

Table 3: Distributional types of the treated species ................................................................................. 31 

Table 4: Main genus delimiting characters and other features of the treated groups ............................. 42 

Table 5: Analysis 1: Species and specimens used in the current study with GenBank accession 

numbers .................................................................................................................................... 295 


numbers .................................................................................................................................... 297 


numbers .................................................................................................................................... 298 

Figures 

I) Species illustrations / distribution maps of the species: 

Fig. 4. / Fig. 5. Chapsa alborosella ..................................................................................... 48 / 49 

Fig. 6. / Fig. 7. Chapsa asteliae ........................................................................................... 50 / 50 

Fig. 8. / Fig. 9. Chapsa astroidea ........................................................................................ 51 / 52 

Fig. 10. / Fig. 11. Chapsa halei ................................................................................................ 53 / 54 

Fig. 12. / Fig. 13. Chapsa indica .............................................................................................. 55 / 56 

Fig. 14. / Fig. 15. Chapsa lamellifera ...................................................................................... 57 / 58 

Fig. 16. / Fig. 17. Chapsa lassae .............................................................................................. 59 / 59 

Fig. 18. / Fig. 19. Chapsa leprieurii ........................................................................................ 61 / 62 

Fig. 20. / Fig. 21. Chapsa leprocarpa ...................................................................................... 63 / 64 

Fig. 22. / Fig. 23. Chapsa lordhowensis .................................................................................. 65 / 66 

Fig. 24. / Fig. 25. Chapsa megalophthalma ............................................................................ 67 / 68


Fig. 26. / Fig. 27. Chapsa magaphlyctidioides ........................................................................ 70 / 70 

Fig. 28. / Fig. 29. Chapsa minor .............................................................................................. 71 / 72 

Fig. 30. / Fig. 31. Chapsa niveocarpa .................................................................................... 73 / 74 

Fig. 32. / Fig. 33. Chapsa phlyctidioides ................................................................................. 75 / 76 

Fig. 34. / Fig. 35. Chapsa platycarpa ...................................................................................... 77 / 78 

Fig. 36. / Fig. 37. Chapsa pulchra ........................................................................................... 79 / 80 

Fig. 38. / Fig. 39. Chapsa subpatens ........................................................................................ 81 / 82 

Fig. 40. / Fig. 41. Chapsa tibellii ............................................................................................. 83 / 84 

Fig. 42. / Fig. 43. Chroodiscus australiensis ........................................................................... 86 / 87 

Fig. 44. / Fig. 45. Chroodiscus parvisporus ............................................................................ 88 / 89 

Fig. 46. / Fig. 47. Fibrillithecis halei ....................................................................................... 91 / 92 

Fig. 48. / Fig. 49. Leptotrema wightii ...................................................................................... 94 / 95 

Fig. 50. / Fig. 51. Leucodecton albidulum ............................................................................. 99 / 100 

Fig. 52. / Fig. 53. Leucodecton compunctelum .................................................................... 102 / 103 

Fig. 54. / Fig. 55. Leucodecton glaucescens ........................................................................ 104 / 105 

Fig. 56. / Fig. 57. Leucodecton occultum ............................................................................. 107 / 108 

Fig. 58. / Fig. 59. Leucodecton subcompunctum ................................................................. 109 / 110 

Fig. 60. / Fig. 61. Melanotopelia rugosa ............................................................................. 112 / 113 

Fig. 62. / Fig. 63. Myriotrema album ................................................................................... 117 / 118 

Fig. 64. / Fig. 65. Myriotrema clandestinum ....................................................................... 120 / 121 

Fig. 66. / Fig. 67. Myriotrema desquamans ......................................................................... 123 / 123 

Fig. 68. / Fig. 69. Myriotrema eminens ............................................................................... 125 / 125 

Fig. 70. / Fig. 71. Myriotrema frustillatum .......................................................................... 126 / 127 

Fig. 72. / Fig. 73. Myriotrema glaucophaenum ................................................................... 128 / 129 

Fig. 75. / Fig. 74. Myriotrema microporum ......................................................................... 131 / 132 

Fig. 76. / Fig. 77. Myriotrema myrioporum ......................................................................... 134 / 134 

Fig. 78. / Fig. 79. Myriotrema olivaceum ............................................................................ 136 / 137 

Fig. 80. / Fig. 81. Myriotrema phaeosporum ....................................................................... 138 / 139 

Fig. 82. / Fig. 83. Myriotrema polytretum ........................................................................... 141 / 142 

Fig. 84. / Fig. 85. Myriotrema protoalbum .......................................................................... 143 / 144 

Fig. 86. / Fig. 87. Myriotrema rugiferum ............................................................................. 145 / 146 

Fig. 88. / Fig. 89. Myriotrema subconforme ........................................................................ 148 / 149 

Fig. 90. / Fig. 91. Myriotrema temperatum ......................................................................... 150 / 151 

Fig. 92. / Fig. 93. Myriotrema trypaneoides ........................................................................ 153 / 154 

Fig. 94. / Fig. 95. Myriotrema viridialbum .......................................................................... 155 / 156 

Fig. 96. / Fig. 97. Nadvornikia hawaiensis .......................................................................... 157 / 158 

Fig. 98. / Fig. 99. Pseudoramonia richeae .......................................................................... 160 / 161 

Fig. 101. / Fig. 100. Reimnitzia santensis ................................................................................ 163 / 164 

Fig. 102. / Fig. 103. Thelotrema adjectum ............................................................................... 167 / 168 

Fig. 104. / Fig. 105. Thelotrema alboolivaceum ..................................................................... 169 / 170 

Fig. 106. / Fig. 107. Thelotrema bicavatum ............................................................................. 171 / 172 

Fig. 108. / Fig. 109. Thelotrema bicinctulum ......................................................................... 173 / 174 

Fig. 110. / Fig. 111. Thelotrema capetribulense ..................................................................... 176 / 176 

Fig. 112. / Fig. 113. Thelotrema circumscriptum..................................................................... 177 / 178 

Fig. 114. / Fig. 115. Thelotrema conveniens ........................................................................... 179 / 180 

Fig. 116. / Fig. 117. Thelotrema crassisporum ....................................................................... 182 / 182 

Fig. 118. / Fig. 119. Thelotrema crespoae ............................................................................... 183 / 184 

Fig. 120. / Fig. 121. Thelotrema cupulare ............................................................................... 185 / 186 

Fig. 122. / Fig. 123. Thelotrema cyphelloides ......................................................................... 187 / 188 

Fig. 124. / Fig. 125. Thelotrema defossum .............................................................................. 189 / 190 

Fig. 126. / Fig. 127. Thelotrema diplotrema ............................................................................ 191 / 192 

Fig. 128. / Fig. 129. Thelotrema eungellaense ........................................................................ 194 / 195 

Fig. 130. / Fig. 131. Thelotrema foveolare .............................................................................. 196 / 197 

Fig. 132. / Fig. 133. Thelotrema gallowayanum ..................................................................... 198 / 199 

Fig. 134. / Fig. 135. Thelotrema lacteum ................................................................................. 200 / 201 

Fig. 136. / Fig. 137. Thelotrema lepadinum ............................................................................ 203 / 204


Fig. 138. / Fig. 139. Thelotrema lepadodes ............................................................................. 207 / 208 

Fig. 140. / Fig. 141. Thelotrema leucophthalmum .................................................................. 210 / 211 

Fig. 142. / Fig. 143. Thelotrema monosporum ........................................................................ 212 / 212 

Fig. 144. / Fig. 145. Thelotrema myriocarpum ........................................................................ 214 / 215 

Fig. 146. / Fig. 147. Thelotrema nostalgicum .......................................................................... 216 / 217 

Fig. 148. / Fig. 149. Thelotrema nureliyum ............................................................................. 218 / 219 

Fig. 150. / Fig. 151. Thelotrema oleosum ................................................................................ 221 / 222 

Fig. 152. / Fig. 153. Thelotrema pachysporum ........................................................................ 223 / 224 

Fig. 154. / Fig. 155. Thelotrema polythecium .......................................................................... 225 / 226 

Fig. 156. / Fig. 157. Thelotrema porinaceum .......................................................................... 227 / 228 

Fig. 158. / Fig. 159. Thelotrema porinoides ............................................................................ 231 / 232 

Fig. 160. / Fig. 161. Thelotrema pseudosubtile ....................................................................... 233 / 234 

Fig. 163. / Fig. 162. Thelotrema rugatulum ............................................................................ 236 / 237 

Fig. 164. / Fig. 165. Thelotrema saxatile ................................................................................. 238 / 239 

Fig. 166. / Fig. 167. Thelotrema saxicola ................................................................................ 241 / 241 

Fig. 168. / Fig. 169. Thelotrema subadjectum ......................................................................... 243 / 243 

Fig. 170. / Fig. 171. Thelotrema subtile ................................................................................... 245 / 246 

Fig. 172. / Fig. 173. Thelotrema suecicum .............................................................................. 247 / 248 

Fig. 174. / Fig. 175. Thelotrema thesaurum ............................................................................ 250 / 251 

Fig. 176. / Fig. 177. Thelotrema triseptatum ........................................................................... 252 / 252 

Fig. 178. / - Thelotrema sp. I ............................................................................................... 253 

Fig. 179. / - Thelotrema sp. II .............................................................................................. 254 

Fig. 180. / - Thelotrema sp. III ............................................................................................. 255 

Fig. 181. / Fig. 182. Topeliopsis acutispora ............................................................................ 258 / 259 

Fig. 183. / Fig. 184. Topeliopsis azorica ................................................................................. 260 / 261 

Fig. 185. / Fig. 186. Topeliopsis darlingtonii .......................................................................... 262 / 263 

Fig. 187. / Fig. 188. Topeliopsis decorticans .......................................................................... 264 / 265 

Fig. 189. / Fig. 190. Topeliopsis elixii ..................................................................................... 267 / 268 

Fig. 191. / Fig. 192. Topeliopsis kantvilasii ............................................................................ 269 / 270 

Fig. 193. / Fig. 194. Topeliopsis laceratula ............................................................................. 271 / 272 

Fig. 195. / Fig. 196. Topeliopsis muscigena ............................................................................ 273 / 274 

Fig. 197. / Fig. 198. Topeliopsis pseudoexanthismocarpa ...................................................... 276 / 276 

Fig. 199. / Fig. 200. Topeliopsis subdenticulata ......................................................................278 / 279 

Fig. 201. / Fig. 202. Topeliopsis tasmanica ............................................................................. 280 / 281 

Fig. 203. / Fig. 204. “Thelotrema” guadeloupensis ................................................................ 283 / 284 

Fig. 205. / Fig. 206. “Thelotrema” parvizebrinum .................................................................. 285 / 286 

Fig. 207. / Fig. 208. “Leptotrema” schizoloma ....................................................................... 287 / 288 

Fig. 209. / Fig. 210. “Thelotrema” subzebrinum ..................................................................... 290 / 291 

Fig. 211. / Fig. 212. “Thelotrema” zebrinum .......................................................................... 292 / 293 

II) Other figures: 

Fig. 1. Main biogeographical zones in Australia ............................................................................. 18 

Fig. 2. Distribution of rainforests in Australia ................................................................................. 19 

Fig. 3. Combined Australian distribution map of all treated species .............................................. 30 

Fig. 213. Phylogeny of Graphidaceae s. lat. as inferred from a two gene-partition analysis ........... 302 

Fig. 214. Phylogeny of Topeliopsis and allied genera of thelotremataceaen Graphidaceae as 

inferred from a nu LSU rDNA data set .............................................................................. 304 

Fig. 215. Phylogeny of Thelotrema and Chapsa (outgroup) as inferred from a mt SSU rDNA data 

set ......................................................................................................................................... 306

0. Deutschsprachige Zusammenfassung 9 

0. Deutschsprachige Zusammenfassung: Taxonomische Studien von 

Mitgliedern der thelotrematoiden Ostroplaes (lichenisierte Ascomycota) 

in Australien 

Die Ordnung Ostropales (Ascomycota) besteht aus sechs Familien und ist durch 

gelatinisierte, hemiangiocarpe Fruchtkörper und funktionell unitunikate Asci gekennzeichnet. 

Wie in dieser Studie gezeigt wird, bilden zwei der Familien, die Graphidaceae und die 

Thelotremataceae, keine einheitlichen monophyletischen Gruppen. Als nomenklatorische 

Konsequenz werden daher die Thelotremataceae mit den Graphidaceae synonymisiert. Die 

ehemalige Familie Thelotremataceae (im Folgenden thelotrematoide Flechten genannt) bildet 

eine der größten Gruppen innerhalb der Ostropales, mit über 1000 beschriebenen Namen 

(Frisch, 2006; Hale, 1981), von denen zur Zeit über 500 als Arten akzeptiert sind (ebd.). 

Diese Arten sind verteilt auf derzeit 21 akzeptierte Gattungen und drei zusätzliche, 

informelle Artengruppen: Acanthotrema, Ampliotrema, Chapsa, Chroodiscus, Diploschistes, 

Fibrillithecis, Gyrotrema, Ingvariella, Leptotrema, Leptotrema schizoloma-Gruppe, 

Leucodecton, Melanotopelia, Melanotrema, Myriotrema, Nadvornikia, Ocellularia, 

Ocellularia clandestina-gruppe, Pseudoramonia, Redingeria, Reimnitzia, Stegobolus, 

Thelotrema, Thelotrema glaucopallens-Gruppe und Topeliopsis. Es handelt sich um 

krustenbildende, vornehmlich lichenisierte Ascomycota, die in der Mehrzahl 

rindenbewohnend sind, und durch einen meist trentepohlioiden Fotobionten (außer 

Diploschistes) und rundliche, eingesenkte bis aufstrebende Peri- oder Apothezien mit 

unverzweigten Paraphysen und distoseptierten Ascosporen charakterisiert sind. Sie bilden 

eine bedeutende Komponente in der Biodiversität tropischer bis subtropischer 

Regenwaldökosysteme, sind aber auch in trockeneren Habitaten und in temperierten Zonen zu 

finden. 

Kenntnisse über Taxonomie, Systematik, Biogeografie und Ökologie in dieser Gruppe sind 

jedoch nur ansatzweise vorhanden, und bis in die jüngste Vergangenheit hinein sind keine 

bedeutenderen systematischen Untersuchungen durchgeführt worden. Die Gründe hierfür 

liegen traditionell zum einen - wie bei nahezu allen regenwaldbewohnenden Individuen - am 

erschwerten Probenzugang und der Problematik der oft kaum erreichbaren Habitate. Zumal 

die Verterter der hier behandelten Gruppe vorzugweise in höheren Bereichen bzw. in der 

Baumkronenzone der Regenwälder zu finden sind. Zum anderen liegen die unterscheidungskritischen 

Merkmale, oft bis in die Familien-Ebene reichend, im mikroskopischen Bereich. 

Mit den unzureichenden optisch-technischen Methoden der historischen Flechtenforschung 

war jedoch eine weitere Differenzierung der Taxa, bzw. die Untersuchung von 

Mikrostrukturen nahezu unmöglich. Zusätzlich spielt in den moderneren taxonomischen und 

systematischen Untersuchungen die Sekundärstoffchemie eine bedeutende Rolle für die 

Unterscheidung verschiedener Gruppen. Die Existenz dieser Substanzen wurde zwar bereits 

in der zweiten Hälfte des 19. Jhs. erkannt, jedoch waren auch hier die technischen 

Analysemöglichkeiten stark eingeschränkt. 

Abgesehen von Redingers Abhandlung (1936) der brasilianischen Vertreter der 

thelotrematoiden Flechten, beschränken sich die Veröffentlichungen bis ins ausgehende 20. 

Jh. in erster Linie auf floristische Untersuchungen, und die systematische Einteilung fußte aus 

heutiger Sicht auf recht groben Kriterien. Als bedeutendste Forscher im Bereich der 

thelotrematoiden Flechten in der zweiten Hälfte des 19. Jhs. sind Nylander (z. B. 1858, 1859, 

1862) und Müller Argoviensis (z. B. 1887b, 1890, 1894) zu nennen. Letzterer führte auch ein 

Ascosporen-basiertes Gattungskonzept (1887b) ein, das bis in die 70er Jahre des 20. Jhs. und 

darüber hinaus Anwendung fand (z. B. Hale 1978, Nagarkar & al. 1988). Demnach wurden 

vier Hauptgattungen unterschieden, die sich nach Septierung und Pigmentierung der 

Ascosporen richteten: Thelotrema mit farblosen, mauerförmigen, Ocellularia mit farblosen


querseptierten, Leptotrema mit braunen, mauerförmigen und Phaeotrema mit braunen, 

querseptierten Ascosporen. Dieses Konzept ermöglichte zwar eine einfache und 

vergleichsweise gut differenzierte Einteilung der Arten, war aber in hohem Maße artifiziell. 

Durch Salisbury (1971, 1972a, 1972b, 1978) wurde daher zunächst ein informelles 

Gliederungssystem eingeführt, das wenige Jahre später von Hale (1980, 1981) in ein 

formalistisch gültiges Gattungskonzept übertragen wurde. Hales Konzept, welches mit 

einigen Modifikationen bis heute Anwendung findet, legt nicht mehr die 

Ascosporenmorphologie zugrunde, sondern berücksichtigt in erster Linie Merkmale des 

Fruchtkörperrandes. 

Es wird im vorherigen Abschnitt bereits andeutungsweise ersichtlich, dass die thelotrematoiden 

Taxa, die lange Zeit ein wissenschaftliches Schattendasein fristeten, mit dem 

Zeitalter der modernen Lichenologie zunehmend in den Fokus der Wissenschaft rückten. Seit 

dem ausgehenden 20. Jh. bis in die heutige Zeit gibt es eine Vielzahl meist kleinerer Studien, 

die sich in erster Linie auf einzelne Untergruppen und/oder auf meist kleinere geografische 

Untersuchungsgebiete beschränken (z. B. Guderley & Lumbsch, 1996; Hale, 1974b, 1978b, 

1981; Homchantara & Coppins, 2002; Kalb, 2001; Kantvilas & Vezda, 2000; Lücking, 1992a, 

Lücking & Grube, 2002; Lumbsch, 1985, 1989; Lumbsch & Elix, 2003; Matsumoto, 2000; 

Nagarkar & Hale, 1989; Nagarkar & al., 1986, 1987, 1988; Patwardhan & Kulkarni, 1977b; 

Patwardhan & al., 1985; Purvis & al., 1995; Salisbury, 1972a, b; Sipman, 1992a, b). Als erste 

Studie, die sowohl auf einer ausführlichen systematischen Untersuchung basiert, als auch eine 

detailierte taxonomische Behandlung nahezu aller Gattungen beinhaltet, ist die von A. Frisch 

und seinen Mitarbeitern kürzlich erschienene Monografie zu nennen (Frisch, 2006; Frisch & 

Kalb, 2006; Frisch & al., 2006), die sich vornehmlich auf den afrikanischen Kontinent 

konzentriert. Für den australasischen Raum allerdings ist das Wissen um die thelotrematoiden 

Flechten in hohem Maße bruchstückhaft. Bis auf die Arbeiten von Lumbsch (1989), der die 

Gattung Diploschistes behandelt, Kalb (2001) und Frisch & Kalb (2006a), die vor allem die 

Gattung Topeliopsis in Australien untersuchen, und Kantvilas & Vezda (2000), die sich auf 

verschiedene Gruppen der tasmanischen Vertreter beschränken, sind keine weiteren 

nennenswerten Studien durchgeführt worden. 

Die hier vorliegende Arbeit hat daher zum Ziel, die taxonomischen Verhältnisse des 

größten Teiles dieser Gruppe für den australischen Kontinent zu beleuchten, und stellt eine 

Revision der trentepohlioiden (d. h. ausgenommen der Gattung Diploschistes), nichtcolumellaten 

(d. h. ausgenommen der Gattungen Gyrotrema, Melanotrema, Ocellularia, 

Redingeria und Stegobolus), thelotrematoiden Graphidaceae (d. h. ausgenommen der 

Graphidaceae s. str.) dar. Im Rahmen des Gesamtprojektes sind die columellaten Taxa 

ebenfalls bearbeitet worden, die Ergebnisse dieser Untersuchungen werden jedoch an anderer 

Stelle veröffentlicht. 

Insgesamt wurden ca. 4000 Proben anatomisch, morphologisch und chemisch untersucht. 

Die untersuchten Individuen setzen sich zusammen aus allen erhältlichen australischen 

Typus-Proben, aus großen Teilen auch außer-australischer Typen, aus allen erhältlichen, 

australischen Aufsammlungen verschiedener Herbarien und schließlich aus den selbstgesammelten 

Proben aus Queensland, Neusüdwales und Victoria. Nach Möglichkeit wurden 

die Proben vorzugsweise anhand des Vergleiches mit dem Typusmaterial bestimmt oder, falls 

dies nicht möglich war, nach den Angaben in den Artbeschreibungen der Literatur. Proben die 

auf diese Art und Weise nicht bestimmt werden konnten, sind entweder provisorisch in 

bereits vorhandene Arten aufgenommen, oder aber als neue Art ausgewiesen worden. Im 

Falle einer zu geringen Probenmenge oder aber wenn sich die Aufsammlung in einem 

schlechten Zustand befand, wurden die Proben vorläufigen, informellen Namen zugewiesen.


So konnten insgesamt 103 Arten bestimmt werden, davon 33 als neu für Australien und 20 

als neu für die Wissenschaft. Von den 71 zur Zeit für Australien bekannten Arten (McCarthy, 

2007) konnten nur 50 bestätigt werden. Die Übrigen stellten sich entweder als anderen 

Gattungen zugehörig, als Synonyme anderer Arten oder aber als falsch bestimmt heraus. Von 

den oben aufgeführten 24 Gattungen und Artengruppen kommen die folgenden 13 in 

Australien vor und sind hier behandelt: Chapsa, Chroodiscus, Fibrillithecis, Leptotrema, 

Leptotrema schizoloma-Gruppe, Leucodecton, Melanotopelia, Myriotrema, Nadvornikia, 

Pseudoramonia, Reimnitzia, Thelotrema und Topeliopsis. Außerdem konnten Spezies 

folgender sieben Gattungen aus Australien identifiziert werden, die in dieser Studie jedoch 

nicht behandelt werden: Ampliotrema, Diploschistes, Ingvariella, Melanotrema, Ocellularia, 

Ocellularia clandestina-Gruppe und Stegobolus. 

Wie bereits erwähnt, wird das auch dieser Studie zugrunde liegende, ursprüngliche 

Gattungskonzept (Hale, 1980, 1981; Salisbury, 1971, 1972a, 1972b 1978) zwar heute noch 

allgemein akzeptiert, wurde aber zum großen Teil weiter ausdifferenziert. Neben der schon 

früh in die Thelotremataceae eingegliederten, und in ihrer Umschreibung seither 

weitestgehendst konstanten Gattung Diploschistes (die sich von allen anderen Gattungen 

durch einen trebouxioiden Fotobionten unterscheidet), sind von den drei trentepohlioiden, 

nach Excipulumstruktur unterschiedenen Hauptgattungen (Thelotrema: Excipulum unkarbonisiert, 

mit lateralen Paraphysen; Myriotrema: Excipulum unkarbonisiert, ohne laterale 

Paraphysen; Ocellularia: Excipulum karbonisiert, ohne laterale Paraphysen) im Laufe der 

Zeit weitere Gattungen nach zusätzlichen Merkmalen der Fruchtkörpermorphologie oder 

sonstigen Kennzeichen ausgegliedert worden (z. B. Frisch, 2006; Frisch & Kalb, 2006; 

Guderley & Lumbsch, 1996; Kalb, 2001, 2004; Kantvilas & Vezda, 2000). So werden die 

hinzugekommenen, hier akzeptierten und behandelten Gattungen des weiteren wie folgt 

unterschieden: 

Laterale Paraphysen und ein unkarbonisierter Ascomarand kommen außer in Thelotrema 

noch in Chapsa, Reimnitzia (hier ist die Exsistenz der lateralen Paraphysen allerdings, wie 

gezeigt werden konnte, zweifelhaft) und Topeliopsis vor. Chapsa ist in erster Linie durch die 

Geaster-artigen, oft sehr großen und offenen Fruchtkörper mit aufrechtem oder zurückgebogenem 

Rand unterschieden; Reimnitzia hat ähnlich geformte Ascomata, ist jedoch in 

verschiedenen Punkten der Thallusmorphologie (Auftreten von Isidien, Thallus auffällig dick 

und mit säulenartig angeordneten Kristallen) und durch mehrere Mikromerkmale des 

Hymeniums, der Asci und der Ascosporen abweichend; Topeliopsis ist charakterisiert durch 

urnenförmige, stark zerschlitzte bis schuppige Ascomata mit auffällig dickem, mit dem 

Thallusrand verwachsenem Excipulum und unverdickten, regelmäßigen und deutlich 

parallelen Paraphysen. 

Eine Sonderform, die sonst nur bei Diploschistes bekannt ist, nämlich das gemeinsame 

Auftreten von lateralen Praphysen und deutlicher Karbonisierung des Excipulums, tritt bei 

den Mitgliedern der Leptotrema schizoloma-Gruppe und in den Gattungen Melanotopelia und 

Pseudoramonia auf. Die erstgenannte Gruppe unterscheidet sich durch regenerierende 

Fruchtkörper mit deutlich mehrlagigen und rissigen Rändern. Melanotopelia hat urnenförmige, 

eher ganzrandige, Pseudoramonia deutlich gestielte Ascomata. 

In den übrigen Gattungen, bei denen gemeinschaftlich laterale Paraphysen fehlen, sind 

Nadvornikia und Chroodiscus am deutlichsten zu unterscheiden. Nadvornikia hat als einzige 

Gattung mazaediöse Ascomata, Chroodiscus umfasst ausschließlich blattbewohnende, 

tropische Arten mit Geaster-artigen Fruchtkörpern. Die Unterscheidung der Gattungen 

Fibrillithecis, Leptotrema, Leucodecton, und Myriotrema ist hingegen schwieriger. Ein sehr 

ähnliches, aus prosoplectenchymatösen Hyphen gebildetes Excipulum ist in Fibrillithecis und 

Myriotrema zu finden. In ersterer Gattung weist es jedoch eine ungewöhnliche, deutlich 

fibrillöse Struktur auf, wohingegen die meisten, und eher typischen Vertreter in Myriotrema,


durch kleine, sehr zahlreich auftretende, eingesenkte Fruchtkörper charakterisiert sind. 

Leptotrema und Leucodecton unterscheiden sich vor allem durch ein paraplectenchymatöses 

Excipulum, die Gattung Leptotrema ist außerdem durch unverdickte Ascuswände und eine 

unterschiedliche Ascosporenentwicklung abgegrenzt. 

Auf den DNA-Sequenzdaten basierende phylogenetische Untersuchungsmethoden sind in 

der modernen Systematik im Laufe der letzten Jahre zu einem wichtigen Bestandteil 

geworden. Die Methoden haben sich gerade im Bereich der Mykologie, wo unzureichende 

oder sehr variabel ausgebildete phänotypische Merkmale die Regel sind, als unverzichtbares 

Werkzeug zur Ermittlung von Verwandtschaftsverhältnissen etabliert. Im Bereich der 

Graphidaceae bzw. ihrer thelotrematoiden Untergruppe sind molekulare Studien bisher 

allerdings rar. Neben einigen Analysen, in die jeweils nur wenige Taxa dieser Gruppe 

miteinbezogen wurden (Grube & al., 2004; Kauff & Lutzoni, 2002; Lücking & al., 2004; 

Miadlikowska & al., 2006; Winka & al., 1998), sind hier noch die eher ausführlicheren 

Arbeiten von Frisch & al. (2006), Lumbsch & al. (2004a), Martín & al. (2003) und Staiger & 

al. (2006) zu nennen, die sich im Spezielleren mit den Graphidaceae s. lat. auseinandersetzen. 

Im Rahmen der vorliegenden Studie wurde deshalb ein Versuch unternommen, die 

Phylogenie der Graphidaceae auf verschiedenen Verwandtschaftsebenen mittels molekulargenetischer 

Verfahren zu beleuchten. Hierzu wurden insgesamt drei Analysen durchgeführt, 

unter Zuhilfenahme zweier molekularer Marker, der mitochondrialen, kleinen Untereinheit 

(mtSSU) sowie der nukleären, großen Untereinheit (nuLSU) der ribosomalen DNA (rDNA). 

Das analysierte Material setzte sich dabei zu großen Teilen aus selbst gesammelten Proben 

der Australienexkursionen sowie aus Frischmaterial anderer Aufsammlungen zusammen. 

Zusätzlich wurden Sequenzdaten benutzt, die in der Internetdatenbank GenBank (USA 

National Center for Biotechnology Information, NCBI) erhältlich waren. 

Das vornehmliche Ziel der ersten, mit einem kombinierten Datensatz beider Gene 

(mtSSU/nuLSU rDNA) operierenden Analyse war es, die Monophylie der Thelotremataceae 

zu testen. Dazu wurden die Sequenzdaten von insgesamt 105 Ascomyceten verglichen. Die 

nahe Verwandtschaft der Thelotremataceae und Graphidaceae, traditionell einzig unterschieden 

durch die Fruchtkörperform (rundlich in Thelotremataceae vs. länglich in 

Graphidaceae), wurde zwar schon in früheren Untersuchungen erkannt und diskutiert (z. B. 

Frisch, 2006; Staiger, 2002; Staiger & al., 2006), genaue Umschreibungen beider Familien 

fehlten jedoch bisher. Das Ziel der beiden anderen Analysen, die jeweils mit einem 

Einzeldatensatz (nuLSU bzw. mtSSU) durchgeführt wurden, war die detailliertere 

Untersuchung der Gattungen Topeliopsis bzw. Thelotrema. Für die nuLSU-Analyse wurden 

dabei 40 Individuen aus 38 verschiedenen Arten der Gattungen Acanthotrema, Chapsa, 

Chroodiscus, Diploschistes, Melanotopelia, Myriotrema, Ocellularia, Thelotrema und 

Topeliopsis getestet. Für die mtSSU-Analyse wurden 50 Individuen aus 25 Arten der 

Gattungen Chapsa und Thelotrema untersucht. 

So konnte gezeigt werden, dass die beiden vormals getrennten Familien Thelotremataceae 

und Graphidaceae phylogenetisch keine einheitlichen Gruppen bilden und ihre bisherige 

systematische Einteilung nicht den natürlichen Verhältnissen entspricht. Darüberhinaus 

konnte gezeigt werden, dass mit Ausnahme der Diploschistes, keine der in die Analysen 

einbezogenen Gattungen in ihrer bisherigen Umschreibung Monophylie aufweisen kann. Es 

wird daher darauf hingewiesen, dass das bisher gebräuchliche Gattungskonzept nach Hale 

(1980, 1981) bzw. Frisch (2006) und Frisch & Kalb (2006) in seiner jetzigen Form weiterer 

Überarbeitungen bedarf. Als nomenklatorische Konsequenz der molekularen Analysen wird 

neben der Synonymisierung der Familien Thelotremataceae und Graphidaceae, die 

Aufstellung einer neuen Gattung Melanotopelia vorgeschlagen.

1. Introduction 13 

1. Introduction 

Over 20% of all known species of fungi are lichenized, i.e. form stable, self-supporting, 

symbiotic associations with photosynthetic partners such as green algae and cyanobacteria. 

Since lichens predominantly belong to the euascomycetes (c. 98%), which results in almost 

half of the taxa in Ascomycota being lichenized, this form of symbiosis is one of the most 

important lifestyles in fungi. Lichens occur worldwide and in all habitats from the tropics to 

the polar regions. They play important ecological roles, including fixation of atmospheric 

nitrogen, stabilizing soil surfaces in semi-arid regions, providing an environment for small 

arthropods or making rocky surfaces inhabitable for other organisms by deteriorating them. 

Despite their importance for a variety of ecosystems, the abundance of lichen species, and 

their omnipresence in the natural landscape, our understanding of biodiversity and 

phylogenetic relationships is surprisingly poor. This is mainly due to the lack of information 

on numerous species, but particularly tropical, crustose groups that exhibit few morphological 

characters are notoriously understudied. 

Within the phylum Ascomycota the order Ostropales currently consists of six families. It is 

characterized by strongly gelatinized ascomata with a hemiangiocarpous ontogeny and 

functionally unitunicate asci. The Thelotremataceae is one of the largest families in 

Ostropales and comprises more than 1000 described names (Frisch, 2006; Hale, 1981), of 

which more than 500 are currently accepted species (ibid.). These usually corticolous, 

crustose lichens have a mostly trentepohlioid photobiont (except in Diploschistes), and are 

further characterized by rounded, immersed to emergent, peri- to apothecioid ascomata with 

unbranched paraphyses and predominantly distoseptate ascospores. They form an important 

component in the biodiversity of subtropical and tropical rainforest ecosystems, but are also 

found in drier habitats and in temperate regions. However, the taxonomic, systematic, 

biogeographical and ecological knowledge of this group is rudimentary, and until most 

recently, no major systematic treatments were available. 

Except for Redinger (1936), who studied the thelotremataceaen Graphidaceae of Brazil, up 

until the end of the 20 th century, the vast majority of publications dealing with this group were 

restricted to floristic treatments and often not more than simple species lists. With the entering 

of modern lichenology, the Thelotremataceae gained more and more attention and several 

studies, predominantly limited to taxonomical approaches of smaller geographical regions 

and/or focused on particular sub-groups were done (e.g. Guderley & Lumbsch, 1996; Hale, 

1974b, 1978b, 1981; Homchantara & Coppins, 2002; Kalb, 2001; Kantvilas & Vezda, 2000; 

Lücking, 1992a, Lücking & Grube, 2002; Lumbsch, 1985, 1989; Lumbsch & Elix, 2003; 

Matsumoto, 2000; Nagarkar & Hale, 1989; Nagarkar & al., 1986, 1987, 1988; Patwardhan & 

Kulkarni, 1977b; Patwardhan & al., 1985; Purvis & al., 1995; Salisbury, 1972a, b; Sipman, 

1992a, b). A first more detailed monograph including the majority of genera and also dealing 

with systematic aspects, was presented by Frisch and his co-workers (segmented in three 

papers: Frisch, 2006; Frisch & Kalb, 2006; Frisch & al. 2006) predominantly addressing the 

thelotremoid lichen biota of Africa. Concerning the research on the Australian continent, 

besides Lumbsch (1989), who treated the genus Diploschistes, Kalb (2001) and Frisch & Kalb 

(2006a), predominantly addressing the genus Topeliosis, and the studies of Kantvilas & 

Vezda (2000) focusing on Tasmania, thus far our knowledge of Thelotremataceae have been 

highly fragmentary. 

The present study is aiming at a revision of trentepohlioid (i.e. excluding the genus 

Diploschistes), ecolumellate (i.e. excluding the genera Gyrotrema, Melanotrema, Ocellularia, 

Redingeria and Stegobolus) thelotrematoid lichens of Australia. The results of the revision of 

columellate taxa for the continent will be presented elsewhere. The study is based on the 

morphological, anatomical and chemical examinations of a total of c. 4000 specimens. These

1. Introduction 14 

specimens include all available type material from Australia (and in large parts types from 

other continents), all accessible herbaria collections of this group from Australia and the 

material collected during two field trips to Queensland, New South Wales and Victoria. 

Where possible, the specimens were assigned to existing taxa, either (preferably) by 

comparison with the type specimen or, if the specific type was unavailable, by verification of 

the data provided in literature. Collections that could not be placed in described species were 

either provisorically included to existing species or new species were introduced. 

Currently c. 20 genera are accepted for Thelotremataceae (Eriksson, 2006; Frisch, 2006), 

16 of which are known to occur in Australia (McCarthy, 2007). The circumscription of the 

genera, however, is controversial. In the traditional classification, the ascospore septation and 

coloration was used schematically to distinguish genera (Müller Argoviensis, 1887b). Since 

the work of Hale (1980, 1981) and Salisbury (1971, 1972a, 1972b 1978), the genera in the 

family are delimited mainly by the structure and pigmentation of the ascomata margin. These 

characters, however, have so far scarcely been tested by molecular data. To approve that the 

inferred phylogenies represent the evolution of the organisms, it was additionally attempted to 

further illuminate the phylogenetic relationships within the family applying molecular 

methods. Therefore, a total of three analyses were carried out employing datasets of two 

different molecular markers from the nuclear and mitochondrial genome.

2. Taxonomic part 15 

2. Taxonomic part 

2. 1. Historical overview 

In traditional lichenology taxonomic concepts have often been subjective and highly 

inconsistent between researchers. Taxonomic groups were circumscribed based on easily 

recognizable features, and even more so in early times of lichen research, when modern 

optical devices were not available. Further, accessible collections were largely restricted to 

northern-hemisphere biotas, while the sampling of tropical regions or other inaccessible parts 

of the world remained poor. Hence, it is not surprising that Graphidaceae, a family of 

predominantly subtropical to tropical taxa that chiefly occur in the canopy of primary 

rainforests and which are often distinguished by micro-morphological characters is still 

poorly understood. With the recently developed tools of molecular phylogeny and a broader 

sampling of these individuals, researchers are gaining a better understanding of the natural 

relationships in this group (e.g., Frisch & al., 2006; Staiger & al., 2006). 

Thus, supported by molecular analysis it could be shown (part 3) that Thelotremataceae 

and Graphidaceae, traditionally separated by their ascoma shape (orbicular vs. lirelliform), 

form several lineages within one strongly supported monophyletic group. Consequently 

Thelotremataceae is regarded synonymous to Graphidaceae. In the following, the focus is set 

on the former family Thelotremataceae, which is thus referred to as thelotremataceaen 

Graphidaceae or thelotrematoid lichens, respectively. I will briefly give an overview of the 

historic development of generic concepts and delimitations within this group. For a more 

detailed treatment dealing with the history of systematics in thelotremataceaen Graphidaceae 

see Frisch (2006), for an historical overview of the developments in Graphidaceae s. str. see 

Nakanishi (1966) and Staiger (2002). 

The family Thelotremataceae was described by Stizenberger (1862) referring to Lecanorei 

subtribe Thelotremei (Nylander, 1861a), although a subtribe Thelotremee was already 

introduced almost ten years earlier by Trevisan (1853a). The first described species in 

thelotremataceaen Graphidaceae was Lichen lepadinus Ach. (Acharius, 1798), which 

subsequently was transferred to its own genus Thelotrema (Acharius, 1803). After several 

additions (Acharius, 1804, 1812a), Thelotrema included seven species (T. cavatum, 

T. discoideum, T. fumosum, T. henatomma, T. obturatum, T. terebratum, T. urceolare) and 

two varieties of T. lepadinum, characterized by a crustose thallus and thelotremoid ascomata. 

Other thelotrematoid taxa were grouped in numerous genera, including Porina, Pyrenula, and 

Urceolaria. With the inclusion of microscopical characters, the classification was redefined 

by Fée (1824, 1825, 1837) and Meyer (1925). Fée introduced two new genera (Ascidium, 

Myriotrema) and accepted Thelotrema (restricted to tropical species) and Volvaria 

(introduced by De Candolle, in Lamarck & De Candolle, 1805) in which he grouped 

T. lepadinum. Meyer did not accept the genus Thelotrema but introduced three new genera 

Ocellularia (later conserved against Ascidium), Anthrocarpum (=Thelotrema) and Porophora 

(nom. superfl. pro Ascidium). 

With increasing appreciation of ascospore characters in the course of the 19 th century the 

recently resurrected genera (Frisch, 2006) Leptotrema (Montagne & Van Den Bosch, 1855), 

Stegobolus (Montagne, 1845), Chapsa and Leucodecton (Massalongo, 1860) were introduced. 

Nylander (1862), however, did only accept two genera, Ascidium and Thelotrema, the latter 

consisting of four informal groups based on ascospore characters. Müller Argoviensis 

(1887b), finally developed a generic concept, solely based on ascospore characters, which 

was accepted until the 1970s (e.g., Hale, 1978; Nagarkar & al., 1988), with four major genera: 

Phaeotrema (ascospores brown, transversally septate), Leptotrema (ascospores brown, 

muriform), Ocellularia (ascospores hyaline, transversally septate) and Thelotrema


(ascospores hyaline, muriform). Three years later, Müller Argoviensis (1890) added 

Chroodiscus for foliicolous taxa. 

This concept was recognized as artificial by many workers, but it was not until the end of 

the 20 th century that attempts were made towards a more natural classification. First Salisbury 

(1971, 1972a, 1972b 1978) proposed to abandon ascospore characters for the generic 

delimitation. He suggested to merge all species into a single genus Thelotrema, with three 

subgenera distinguished by the morphology of the ascoma margin, each divided into different 

species groups characterized by further ascomata features: Thelotrema sect. Ascidium 

(including the T. discolor-, T. cavatum- and T. discoideum-group) - carbonized with absent 

lateral paraphyses; Thelotrema sect. Myriotrema (including the T. compunctum- and the T. 

bahianum-group) – non-carbonized with absent lateral paraphyses; Thelotrema sect. 

Thelotrema (including the T. lepadinum- and the T. platycarpum-group) - non-carbonized 

with present lateral paraphyses. Subsequently, Hale (1980, 1981) implemented Salisbury’s 

concept with minor modifications and introduced three major genera equivalent to Salisbury’s 

subgenera: Myriotrema (=T. sect. Myriotrema), Ocellularia (=T. sect. Ascidium) and 

Thelotrema (=T. sect. Thelotrema). Ever since, Hale’s classification was broadly accepted 

(e.g., David & Hawksworth, 1995; Sipman, 1993, 1994; Matsumoto, 2000; Homchantara & 

Coppins, 2002; Frisch, 2006) and is still in use, although it was also soon realized that the 

distinction in three large genera was too coarse and did not mirror phylogenetic relationships. 

However, some authors (Poelt & Vezda, 1981; Purvis & al., 1995) rejected Hale’s 

classification and suggested to place all core genera in one large genus Thelotrema following 

Salisbury’s proposal. On the other hand, numerous smaller homogeneous groups were divided 

and introduced as separate genera in subsequent years: Ampliotrema (Kalb, 2004), 

Chroodiscus (Kantvilas & Vezda, 2000; Lücking, 1992; Lücking & Kalb, 2000; Lumbsch & 

Vezda, 1990), Ingvariella (Guderley & Lumbsch, 1996), Nadvornikia (Tibell, 1984), 

Pseudoramonia (Kantvilas & Vezda, 2000), Reimnitzia (Kalb, 2001), Topeliopsis (Kantvilas 

& Vezda, 2000; Kalb, 2001). 

Recently, Frisch (2006) and Frisch & Kalb (2006) described five new genera 

(Acanthotrema, Fibrillithecis, Gyrotrema, Melanotrema, Redingeria) and resurrected four 

formerly described genera (Chapsa, Leptotrema, Leucodecton, Stegobolus). Meanwhile, other 

genera formerly included were transferred to other families, e.g. Conotrema to Stictidaceae 

(Eriksson & al., 2003), Gyrostomum to Graphidaceae (Staiger, 2002), Ramonia to 

Gyalectaceae (Eriksson & al., 2003), Tremotylium (=Minksia) to Roccellaceae (Makhija & 

Patwardhan, 1995). The systematic position of Phaeotrema is still uncertain, according to 

Salisbury (1978) its type species Pyrenula subfarinosa is a non-lichenized fungus. In contrast 

to the above mentioned three main genera, Diploschistes, an additional, species-rich genus 

included in Thelotremataceae, forms a homogeneous species-group that is also well supported 

by molecular data (Lumbsch & al., 2004a; Frisch & al., 2006). It was placed in its own family 

(Diploschistaceae) by Zahlbruckner (1905), but has been placed in Thelotremataceae for a 

long time (Gilenstam, 1969). 

2. 2. Taxonomic studies and collections of trentepohlioid thelotremataceaen 

Graphidaceae in Australia 

Until recently all contributions to the knowledge of thelotrematoid lichens in Australia 

were restricted to floristic studies. The majority of known species from this continent are 

based on collections made by early naturalists in the late 19 th century. Amongst the most 

important collectors of this time were F. M. Bailey (1827-1915), an appointed colonial 

botanist and curator at the Queensland Museum (Brisbane); the New Zealand botanist and 

passionate lichenologist C. Knight (1808-1891), who collected in New Zealand, but also in 

Queensland and New South Wales; the educationist and scientist J. Shirley (1849-1922), who


conducted field trips throughout Queensland; the naturalist and plant collector W. A. Sayer 

(fl. 1886-1897) who was an official participant of expeditions to Pacific northern Queensland; 

and the Presbyterian minister and amateur botanist F. R. M. Wilson (1832-1903), who is 

regarded the pioneer in Australian lichenology. He mainly collected in Victoria, but also in 

New South Wales and southern Queensland. 

The first record of Australian thelotrematoid lichens was made by Nylander and 

Krempelhuber (in Nylander, 1864). They introduced Thelotrema bicavatum and T. lacteum 

based on collections of Hochstetter from an unknown locality in Australia. Subsequently, 

Stirton (1881) described T. profundum [=Ocellularia profunda] and Endocarpon baileyi 

[=Leptotrema wightii] based on Bailey collections. The most active worker on Australian 

lichens in the late 19 th century, however, was Müller Argoviensis (1882, 1887a, b, 1888a, 

1891a, b, 1892, 1893a, b, 1895d), who described 36 species and one variety for Australia. The 

new taxa were mostly based on collections from Sayer, C. Knight, Shirley, Bailey and Wilson 

from the Brisbane and Cairns area (Queensland) and other locations in New South Wales and 

Victoria. His contemporary C. Knight described five species for Australia based on his own, 

Bailey’s and Shirley’s collections from southern Queensland. Another species from southern 

Queensland was described by Wilson (1893), based on his own collection: O. cricota [=T. 

lacteum]. 

Besides two publications by Jatta (1911), who introduced the first Tasmanian taxa of 

trentepohlioid thelotrematoid lichens, and Räsänen (1949) who described two new species 

and three varieties in the course of a review of Wilson collections in H, for most of the 20 th 

century, however, thelotremataceaen Graphidaceae in Australia have been neglected. It was 

until recent times before this group regained scientific attention, and several new taxa were 

described: Chroodiscus australiensis (Lumbsch & Vezda, 1990); C. asteliae [=Chapsa], C. 

lamelliferus [=Chapsa lamellifera], C. australis ssp. tasmanicus [=Chapsa tasmanica], C. 

minor [=Chapsa minor], Topeliopsis muscicola [=T. muscigena], Pseudoramonia richeae, 

Topeliopsis rugosa [=Melanotopelia] (Kantvilas & Vezda, 2000); T. acutispora, T. corticola 

[=T. decorticans], T. vezdae [=T. subdenticulata] (Kalb, 2001b); T. darlingtonii, T. elixii 

(Frisch & Kalb, 2006a); Stegobolus carneopustulatus (Frisch & Kalb, 2006b); Thelotrema 

eungellaense, T. gallowayanum (Mangold & al., 2007a); and Ocellularia kalbii (Mangold & 

al., 2007b). 

For the present treatment I studied a large number of samples collected by various 

researchers collected within the last c. 40 years in addition to material gained in two field trips 

that were particularly focused on thelotrematoid lichens. The largest collections include those 

by J. Elix, J. Hafellner, M. E. Hale, K. Kalb, G. Kantvilas, H. Streimann and L. Tibell. 

2. 3. Geography of Australia 

Australia is the world’s smallest continent (7,741,220 km 2 ) and is situated between the 

Indian and the Pacific Oceans (c. 10-43º S [including Tasmania] and 113-153º E). The 

Australian continent comprises a variety of biogeographical regions, ranging from deserts to 

tropical rainforests and seacoasts to alpine reaches of up to 2,228 m (Mt. Kosciuszko, New 

South Wales). In this chapter, I will provide a brief overview of the climate and the vegetation 

forms of Australia with special emphasis on the distribution of the here treated species group. 

A more extended summary of the Australian Vegetation is given by Beadle (1981), Specht 

(1970, 1981), Specht & Specht (1999) and also Walter & Breckle (1991). Lichenological 

aspects of this topic are discussed by Stevens (1987) and Lumbsch (1994), in Barlow (1981) 

the historical developments of the flora in Australia are treated. A detailed presentation of the 

ecological relationships of the Tasmanian lichen biota is provided by Kantvilas & al. 

(Kantvilas, 1988, 1990; Kantvilas & Jarman, 1988, 1991; Kantvilas & Minchin, 1989).


Fig. 1. Main biogeographical zones in Australia (for explanation see text). 

Most of Australia is dominated by arid climates with dry grass- or shrublands or dry 

woodlands or more rarely dry forests (fig. 1: zone 1). Areas with higher precipitation are 

restricted to the oceanic regions of northern, eastern and parts of southern Australia. These 

regions are distinguished in the tropical and subtropical northern and northeastern sections 

(zones 2+3), and the temperate southeastern and southwestern sections (zones 4+5). The 

south of Western Australia as well as parts of South Australia and Victoria (zone 5) have a 

Mediterranean climate with a winter rain season and are dominated by dry sclerophyll forests 

(Western Australia), Mallee woodlands and -shrubs or, in drier areas, Chenopod shrublands. 

A temperate climate with a more uniform distribution of precipitation periods is found in 

south-eastern Australia, including Tasmania, most of Victoria and eastern New South Wales 

(zone 4), where cool- to warm-temperate rainforests and wet or dry sclerophyll forests occur. 

However, the native vegetation forms of this region have been highly transformed by 

anthropogenic factors, and major primary extends have been cleared. An overview of all 

currently existing rainforest areas is given in fig. 2. 

Remnants of subtropical and tropical low- and highland rainforests (fig. 2) and in drier 

regions several forms of Eucalypt forest communities as well as mangroves are found along 

the northeastern Pacific coast of Queensland, to the west limited by the Great Dividing 

Ranges (zone 3). This comparatively narrow strip includes the humid regions in Australia, 

with predominantly monsoonal climate and peak precipitations during the rain seasons in 

summer and winter. Only in the wider Cairns area in northern Queensland higher rainfalls 

persist throughout the year. Most of northern Australia, however, has a tropical monsoonal 

climate with higher rainfalls restricted to the summer months (zone 2). These regions are 

dominated by savannah with gallery forests, but also smaller areas of rainforests and more


humid Eucalypt forests as well as mangroves along the east coast of Cape York (Queensland) 

and north Northern Territory. 

Fig. 2. Distribution of rainforests in Australia (black dots). 

2. 4. Materials and Methods 

2. 4. 1. Examined collections 

Specimen and type material of thelotremataceaen Graphidaceae from the following 

herbaria was examined: ABL, B, BCRU, BG, BM, BP, BRI, C, CANB, COLO, DUKE, E, F, 

FH, G, GLAM, GZU, H, HIRO, HO, L, M, MEL, MSU, NSW, NY, PERTH, RAMK, S, 

TNS, TUR, UPS, US, WELT, WIS and the private herbarium of Klaus Kalb (Germany). 

Also, the following herbaria were visited to study and loan collections: C, F, US and WIS. 

Additionally material was collected on two field trips. The first in September and October 

2003 in collaboration with T. Lumbsch, where the focus was set on the biotas of Pacific 

Queensland, the second in February and March 2005, in partial collaboration with J. Elix, 

where collections were made in Victoria, Pacific New South Wales and northern Queensland. 

The specimens were collected using a knife or hammer and chisel, and the material was 

stored in consecutively numbered paper bags where they remained for drying. Specimens on 

certain types of bark were also additionally pressed in a plant press (to avoid the curling of the 

bark). In total, about 4000 specimens were examined for this study.


2. 4. 2. Methods 

All specimens were morphologically, anatomically and chemically examined. Habit 

characters were observed by using a low-magnification dissecting microscope (Zeiss Stemi) 

with magnifications from 10x to 50x. Further, thalli and ascomata were cut using a razor 

blade and a freezing microtome (Leica SM2000R), and examined in water, iodine (Lugol’s 

solution, Merck), potassium hydroxide solution and lactophenol cotton blue with a compound 

microscope (Zeiss Axioskop 2plus) at magnifications from 100x to 630x. Images of the habit, 

ascomata- and thallus-sections and ascospores were taken by using the above-mentioned 

optical devices, a camera-fixture and a digital mirror reflex finder camera (Nikon Coolpix). 

The images were subsequently processed with the computer programs iPhoto and Adobe 

Photoshop. 

Secondary metabolites were identified by examination under UV-light, spot tests, and thin 

layer chromatography. Spot tests were made by applying drops of potassium hydroxide 

solution (K), calcium hypochlorite solution (C) and paraphenylenediamine solved in ethanol 

(PD) directly on the thallus and the ascomata of the specimen. 

Thin layer chromatography (TLC) was carried out after Lumbsch (2002) using acetone 

extractions of preferably substrate-free samples of the lichen thallus and ascomata, that were 

applied on silica plates and developed in two solvent systems: A = toluene/dioxin/glacial 

acetic acid (180:45:5), and B’ = hexane/methyl tert-buthyl ether/formic acid (140:72:18). In 

two steps, the plates were first observed under UV-light, and finally after treatment with 10% 

concentrated sulfuric acid solution and charring at 110º C. The secondary metabolites finally 

were determined on the plates according to their color after charring (with and without UVlight) 

and their position in relation to the reference pure substances (stictic, norstictic and 

psoromic acid and atranorin) and the front of the solvent system (Rf-class). 

An additional molecular examination was carried out for selected specimens of the fresh 

material from the above-mentioned field trips; see part 3 for material and methods. 

2. 4. 3. Citations 

Author names of taxa are cited following Kirk & Ansell (1992). Descriptions of collection 

localities and other data concerning the examined specimens are given as follows: Country, 

State/Province and - according to the provided data - additional administrative units and/or 

further descriptions of the site, date (only if the first mentioned specifications are fragmentary 

or missing), collector(s) name(s) and collecting number (herbarium acronym[s], voucher 

number of the herbarium [if name of collector or collecting number is missing]). 

2. 4. 4. Abbreviations 

B/MCMC: Bayesian analysis using Markov chain Monte Carlo methods 

C: Calcium hypochlorite solution 

Distr.: District 

Hwy.: Highway 

I: Iodine 

Isl.: Island(s) 

K or KOH: Potassium hydroxide solution (10%) 

LSU: Large subunit 

MP: Maximum parsimony 

mt: Mitochondrial 

NP.: National Park 

nu: Nuclear


PD: Paraphenylenediamine solved in ethanol 

PP: Posterior probability 

rd.: Road 

rDNA: Ribosomal DNA 

SF.: State Forest 

SSU: Small subunit 

TLC: Thin layer chromatography 

UV: Ultraviolet light 

!: Type specimen studied 

2. 5. Main characters of trentepohlioid thelotremataceaen Graphidacae 

In the following I will provide a brief introduction of the main characters of thelotrematoid 

Graphidaceae. A more detailed account can be found in chapter 2. 9., where the individual 

genera are discussed. Besides Frisch’s (2006) detailed studies of the morphology and anatomy 

of thelotrematoid lichens, only a few more specific publications are available. A detailed 

treatment of the ascoma ontogeny, in particular of columellate taxa, can be found in 

Redinger's monograph of the Brazilian Thelotremataceae (1936). In Hale (1974a, 1981) the 

focus is set on thallus anatomy with special emphasis of cortex structures. A study dealing 

with pycnidial structures in thelotrematoid lichens was done by Matsumoto & Deguchi 

(1999). Concerning the genus Diploschistes, more detailed information is found in Lumbsch 

(1989) and Lumbsch & al. (1997), where particularly the ascospore ontogeny of the genus is 

discussed. 

2. 5. 1. Thallus 

Morphology 

As in other species of Graphidaceae the thallus of thelotrematoid lichens is crustose. It may 

be hyposubstratic or episubstratic, in a few cases the thallus is partly bulging and ±flaking 

away from the substrate. The thallus size and thickness is highly variable and ranges from a 

few millimeters in diameter (e.g. in Chroodiscus) and entirely hyposubstratic thalli (e.g. 

Thelotrema pachysporum, “Thelotrema” guadeloupensis) to specimens that cover large areas 

of sometimes several decimeters in diameter (e.g. in Myriotrema album or Thelotrema 

porinaceum) and can be up to 1 mm high (e.g. Leptotrema wightii, Leucodecton glaucescens). 

In some taxa, a high intraspecific variability in thallus morphology could be observed due to 

altering habitat factors, such as humidity or density and hardness of the substrate. The thallus 

color varies from bright to dark and may be grayish to greenish or brownish. Especially in 

thin thalli, the thallus color is often influenced by the substrate. 

The thallus surface can vary from shiny to ceraceous to dull or pruinose, and from 

continuous to rugose to distinctly verrucose or verruculose. In some species of various genera 

the surface has a reticulate structure and/or forms a grainy-speckled pattern caused by large 

crystals in the medulla (e.g. Leptotrema wightii, Leucodecton glaucescens Myriotrema 

phaeosporum, Reimnitzia santensis, Thelotrema oleosum). The thalli are either unfissured or 

distinctly fissured to sometimes rimose or areolate. A prothallus is formed by most taxa 

(absent in Chroodiscus), which is predominantly rather indistinct and thin and brown. 

Anatomy 

The thallus anatomy in thelotremataceaen Graphidaceae is variable. Although some 

species are stratified, and upper cortex, algal layer and medulla can be distinguished, most 

taxa show a more diffuse anatomical structure. A thallus cover is either absent, or consists of 

a variably thick layer of weakly conglutinated and loosely arranged hyphae (protocortex). In


some species, a variably thick true cortex of highly conglutinated and ±densely arranged to 

cartilaginous irregular or periclinal hyphae is present. Many species also show intermediates 

of these cortex types, sometimes even within a single specimen. In contrast, Frisch (2006), 

who follows the terminology of Poelt (1989) and Büdel & Scheidegger (1996) summarizes all 

cortex structures under the term ‘phenocortex’ and lists four different types (see there). 

The species treated here all have a trentepohlioid photobiont. Abundance of algal cells 

differs widely between specimens. In some species the algal layer is well-developed and 

continuous but can be also discontinuous or almost absent. In this case photobiont cells are 

scattered throughout the entire thallus. In predominantly hypophloedal specimens, algal cells 

are also found amongst the hyphae that penetrate the substrate layer. The algal layer is often 

interrupted by large oxalate crystals, in some species these crystals are organized in columns. 

The presence of oxalate crystals is also variable. Species that produce secondary metabolites 

may contain additional thalline crystals; the most prominent example is Leptotrema wightii, 

which contains bright red anthraquinone crystals. A distinct medulla layer is rather unusual 

and occurs in epiphloedal taxa with thicker thalli. In Reimnitzia santensis sometimes lower 

cortex-like structures are found in the basal thallus regions. 

Vegetative propagules 

Several forms of vegetative reproduction are known for thelotrematoid lichens, isidia occur 

in two Australian species of Myriotrema and in Reimnitzia (in Pseudoramonia isidia-like 

structures are present, which probably represent immature ascomata). Soralia are known from 

one member of Leucodecton, otherwise soralia as wells as schizodiscs are predominantly 

known for columellate species (not treated here). 

2. 5. 2. Ascomata 

Morphology 

Ascomata in thelotremataceaen Graphidaceae are predominantly hemiangiocarpous, perito 

apothecioid or Geaster-like and open by a single pore, or mazaedious. Sizes of mature 

ascomata range from c. 100 µm to c. 5 mm in diameter. They are predominantly roundish to 

slightly irregular, occur solitary to distinctly fused and their position relative to the thallus 

reaches from deeply immersed to distinctly emergent or stipitate. In some genera, ascomata 

develop in succession (Pseudoramonia, Melanotopelia) or are regenerative (Chapsa, 

Leptrotrema schizoloma-group). A well developed, incurved to recurved, entire to lacerate 

thalline rim is mostly present, in some cases it can become strongly eroded with age. The 

thalline rim is usually concolorous with the thallus or brighter, rarely whitish or 

conspicuously stained. Ascomata discs are flat to concave and often visible from the surface. 

They are predominantly whitish, grayish, brownish, flesh-colored or rarely distinctly reddish, 

and are epruinose to distinctly pruinose. 

The ascoma morphology is an important character in thelotrematoid lichens that delimits 

several genera. The following morphological ascomata types can be distinguished: 

– Geaster-like (chroodiscoid): predominantly large, conspicuous, immersed-erumpent, 

gaping, with distinctly exposed disc and fissured to lacerate or eroded, often layered 

margin; margins erect to recurved or exfoliating and usually proper exciple fused and not 

visible; ascomata regeneration common: Acanthotrema (not treated), Chapsa, Chroodiscus 

and Reimnitzia. 

– mazaedious: large, conspicuous, distinctly emergent with visible mazaedium; nonregenerating: 

Nadvornikia. 

– stipitate: medium-sized, conspicuous, peri- to apothecioid with distinctly stiped base 

and successive growth: Pseudoramonia.


– myriotremoid s. str.: predominantly small and numerous, inconspicuous, immersed, 

apothecioid with non-visible to partly visible disc; proper exciple free to fused, thallus 

margin entire and incurved to erect; non-regenerating: Leptotrema, Leucodecton and 

Myriotrema s. str. (=’M. olivaceum-group’). 

– thelotremoid s. str.: predominantly medium-sized to large, ±conspicuous, emergentsessile, 

apothecioid with partly visible disc; proper exciple visible in large parts, thallus 

margin entire to split, incurved to erect; exciple and thallus margin forming a double-pore; 

non-regenerating: Thelotrema (p. pt.) and Myriotrema s. lat. p. pt. (‘Myriotrema viridialbum-group’). 

– perithecioid-thelotremoid: medium-sized to large, inconspicuous, immersed to 

emergent, perithecioid; apical proper exciple visible, fused or free, thallus margin entire to 

slightly split, incurved; non-regenerating: Thelotrema and Myriotrema s. lat. p. pt. (M. 

desquamans, M. trypaneoides). 

– emergent-perithecioid: large, conspicuous to inconspicuous, ±emergent, perithecioid; 

proper exciple not visible or apically visible, then fused and fibrous, thallus margin 

incurved; non-regenerating: Fibrillithecis and Myriotrema s. lat. p. pt. (M. eminens, M. 

frustillatum), . 

– topeliopsioid: large, conspicuous, emergent-sessile and predominantly subglobose to 

urceolate, peri- to apothecioid, then with partly visible, epruinose disc; proper exciple not 

visible, margin bright, squamulose to pruinose and incurved, sometimes layered, then 

lacerate and exfoliating; non-regenerating or with successive growth: Melanotopelia and 

Topeliopsis. 

– layered-carbonized: small to large, conspicuous to inconspicuous, immersed-erumpent 

to raised, peri- to apothecioid, then with partly visible, epruinose disc; proper exciple 

visible but indistinct, margin split to lacerate and distinctly layered, layers concentrically 

striated, alternating in dark and bright; regenerating: Leptrotrema schizoloma-group. 

Sterile specimens were generally not treated, except for sterile collections of M. 

frustillatum that could be identified by their isidia-type and chemistry, and four other sterile 

collections that were tentatively included in T. bicinctulum. 

Anatomy 

The presence of lateral paraphyses is a unique character that is only known from 

Ostropomycetidae. The presence or absence of these is a generic character in the family. 

Further characters of ascomata anatomy at generic level include the structure of the proper 

exciple and, to some extend, the structure of the paraphyses-tips and the micro-morphology of 

the ascus walls (for a compilation of all genera delimiting characters see also table 4) The 

ascospore morphology, on the other hand, is of major importance for the delimitation of 

species. Further, in several genera (which will be treated elsewhere) a columella is present. 

Proper exciple: The proper exciple can be entirely fused to the thalline rim or be entirely 

free, with several intermediate stages. Its thickness ranges from evanescent or very thin to 

very thick, and it is either formed of paraplectenchymatous or prosoplectenchymatous 

hyphae. Usually it is hyaline or yellow, brown or orange internally and becomes more dark 

colored towards the margins and apices. Several taxa have distinctly dark or slightly 

carbonized proper upper exciple, an entirely distinctly dark or carbonized exciple is present in 

the members of the Leptotrema schizoloma-group, Melanotopelia and Pseudoramonia. The 

proper exciple can include substrate particles and/or is often apically ±covered by granules. 

Many taxa show a ±distinct amyloid reaction, which is mostly located in the lower exciple at 

the intersection to the subhymenium.


Frisch (2006) distinguishes three different exciple types in his revised generic concept, 

which, however, include the presence or absence of lateral paraphyses. Accordingly, in 

Myriotrema and Fibrillithecis an ‘Ocellularia-type’ exciple is present, a ‘Leucodecton-type’ 

exciple is found in Leptotrema and Leucodecton, and a ‘Thelotrema-type’ exciple is 

characteristic for the genera with chroodiscoid ascomata and Thelotrema and Topeliopsis. 

(For further comments and explanations of the exciple-types see under chapter 2.9., but see 

also in Frisch [ibid.]). 

Subhymenium and hypothecium: The subhymenium is predominantly indistinct, 

evanescent to thin, rarely thick, and usually concolorous with the proper exciple, but 

sometimes also conspicuously dark pigmented. In some species with regenerating ascomata, a 

distinct, hyaline, gelatinous I+ purple hypothecial area can be present, which represents newly 

developing hymenial tissue. 

Hymenium: The colorless, non-amyloid hymenium in thelotrematoid lichens is discoid to 

cupular, clear to inspersed, weakly to strongly conglutinated and can be from c. 50 to 400 µm 

high. The paraphyses are simple to slightly branched, c. 1-3 µm thick, straight to ±bent or 

apically curly, parallel to ±interwoven. Sometimes also irregular paraphyses with a distinct 

septation occur. The paraphyses tips are either not thickened or ±distinctly thickened and 

simple to ±irregular. As mentioned above, all of the here treated species lack a columella. 

Nevertheless, columella-like structures often occur in species with strongly fused ascomata 

and resemble excipular tissue. 

Lateral paraphyses: Lateral paraphyses (following Henssen, 1995) (‘periphysoids’ in 

Frisch [2006]) are present or absent. They are mostly clearly separated from the proper 

exciple, but sometimes also basally difficult to distinguish from it. They are predominantly 

clear, very rarely interspersed and range from up to c. 10-50 µm in length. Although the 

presence of lateral paraphyses is usually easy to recognize, in some cases inconspicuous and 

easily overlooked lateral paraphyses occur. 

Epihymenium: An epihymenial layer is variously developed, it is indistinct to very thick, 

predominantly hyaline to more rarely yellowish or brownish, in Chroodiscus australiensis 

conspicuously orange to reddish. It can be egranulose to distinctly granulose, the granules are 

fine to coarse and grayish to brownish. 

Ascus: The asci are 1-, 2-, 4- or 8-spored, non-amyloid, unitunicate and clavate. The ascus 

walls are usually not thickened and a ±distinct tholus with an absent to small, tapered to 

roundish ocular chamber is developed. The tholus mostly becomes, particularly in asci 

bearing large ascospores, ±indistinct in later stages of development. However, several deviant 

forms of asci could be observed and the taxonomical relevance of this feature is unclear. 

Frisch (2006) lists up to five main ascus-types, and for the re-introduced genus Leptotrema he 

considers the ascus morphology as a main delimiting character. In my observations, I could 

find inconsistencies in several taxa, a ‘Leptotrema-type’ ascus (distinct tholus absent, ascus 

walls evenly thickened in younger stages, thin at maturity) could also be found in other 

genera, for example in Chapsa (C. lamellifera), Leucodecton (L. compunctellum), Myriotrema 

(M. frustillatum) or Thelotrema (T. oleosum); a tholus with a large, distinctly tapered ocular 

chamber, reported for Chapsa eitenii and C. zahlbruckneri (Frisch ibid.) also occurs in M. 

protoalbum (see also table 4). 

Ascospores: The ascospores occur uni- to triseriate, rarely quadriseriate, range from 6 up to 

400 µm in length and form 2 up to 55 µm in width. They can be transversely septate or


submuriform to densely eumuriform, with a single to multiple loci; hyaline to distinctly 

brown and non-amyloid to strongly amyloid. The ascospores are mostly straight, but can also 

be distinctly bent, their form can be subglobular, ellipsoid, oblong, clavi- or fusiform, 

bifusiform, bacillar-fusiform, bacillar or cylindrical with roundish to distinctly acute ends. In 

some taxa, particularly in members of Ocellularia, the ascospore ends are conspicuously 

tapered and appendix-like (see also Mangold & al., 2007b). The ascospore wall consists of a 

variably thick endospore, a variably thick epispore and a variably thick or absent perispore (in 

the following also referred to as halo). The epispore marginally determines the shape of the 

ascospore, internally it is delimited by the loci and near the septae it merges into the 

endospore (in the following, the epispore is referred to as cell wall). The marginal epispore 

(also referred to as ‘exospore’) is mostly smooth, but sometimes – often towards the late 

maturity - it can be crenate or ornamented. The form of the endospore is internally determined 

by the loci and only distinct in eumuriform ascospores, in transversely septate and 

submuriform ascospores it internally merges with the septae (in the following the endospore 

is thus only specified in muriform ascospores). The halo (perispore) consists of a gelatinous 

substance that dissolves in KOH, and is usually distinct only in younger stages of the 

ascospore development. It is predominantly continuous, but in some taxa it can also be 

±irregular. Frisch (2006) distinguishes further ascospore structures as a ring-like structure that 

resembles the internal loci wall, a middle lamella along the central septum, and two further 

layers of the ascospore wall, a mesospore and an ‘epispore’, which, according to Frisch (ibid.) 

- who follows the terminology of Janex-Favre (1964) - is used to describe a thin, gelatinous 

coating of the ascospore. Frisch (ibid.) also introduced the expressions of the ‘thin-walled 

type’ and the ‘thick-walled type’ ascospores to distinguish the ascospores of the genera 

Acanthotrema and Chroodiscus (having ‘thin-walled type’ ascospores) from the remaining 

genera. However, this phrasing is somewhat misleading since ‘thick-walled type’ ascospores 

can also have distinctly thin ascopore walls. The septation of the ascospores varies from 

species having distinct and thick septae (particularly in transversely septate ascospores), to 

species with indistinct, thin or absent septation, which can occur in multi-locular ascospores 

at late maturity or in small, submuriform ascospores with irregular loci arrangement. Further, 

the septation can be continuous and regular to discontinuous and irregular. The shape of the 

loci is roundish to angular, and subglobose, oblong, lentiform, cuboid or irregular; the end 

cells (if solitary) are either of the same shape as the remaining loci to more often 

hemispherical or conical. 

Two special forms of ascospores are ascoconidia producing ascospores and the ascospores 

characteristic for the ‘Ocellularia clandestina-group’. The first are found in two species of 

Topeliopsis and will be treated below, the latter are found in Chapsa halei and in a recently 

introduced (Frisch ibid.) sub-group of Ocellularia (not treated here) and have been so far only 

known from there. The ascospores are mainly deviant in having strongly thickened walls that 

are non-amyloid, and angular loci that – in early development – are separated by “pored 

septae” (Frisch, ibid.: 334) and become centrally fused in later maturity. 

2. 5. 3. Conidiomata 

Pycnidia are known for all treated genera (the presence of pycnidia for Chapsa is newly 

reported) except Chroodiscus, Melanotopelia, Nadvornikia, Pseudoramonia and Topeliopsis. 

In two species of the latter genus, however, old ascospores generate ascoconidia (see under 

Topeliopsis for details). The pycnidia are either immersed or in ±strongly raised thallus warts, 

and open by a single, tiny pore, that often is surrounded by a dark area. The conidia are either 

bacilliform, fusiform, oblong, obovate or irregular, and range from 2-8(10) x 0.5-2 µm in size. 

For a more detailed treatment of conidiomata in thelotremataceaen Graphidaceae see Frisch 

(2006).


2. 5. 4. Chemistry 

In the majority (approximately 70%) of the species secondary compounds are present, 

whereas in Graphidaceae s. str., lichen substances occur less frequently (Staiger 2002). The 

majority of compounds are β-orcinol depsidones; in addition, orcinol depsides (chiefly 

occurring in Diploschistes), xanthones, anthraquinones (and other pigments) and several 

unidentified substances are known. 

The taxonomic relevance of secondary compounds in lichenized fungi was controversially 

discussed in numerous publications (e.g., Brodo, 1978; Culberson, 1969, 1986; Culberson & 

Culberson, 1976, 1994; Hawksworth, 1976; Elix, 1993; Elix & al., 1986; Esslinger, 1977, 

1989; Feige & Lumbsch, 1995; Lumbsch, 1998a, b; Rogers, 1989). Particularly at the species 

level, the taxonomic interpretation of chemical differences remains problematic and needs 

further molecular studies. In the present treatment, morphologically indistinguishable 

chemotypes were consistently regarded conspecific and designated to chemical strains. 

In the following, a compilation of detected compounds is provided, grouped in compound 

classes and chemosyndromes (where present). Newly identified substances (Elix, pers. com.) 

or unknown substances are marked with an asterisk and specified in a separate table. For Rfvalues 

and spot characteristics of the known substances see Elix & al. (1995, 2000c) and 

Huneck & Yoshimura (1996). 

β-Orcinol depsidones 

Salazinic acid 

Detected as singular, major compound in Thelotrema circumscriptum and ‘Leptotrema’ 

schizoloma (strain II). 

Stictic acid chemosyndrome 

Includes variable concentrations of α -acetylconstictic, α-acetylhypoconstictic, 

consalazinic, constictic, cryptostictic, hypoconstictic, hyposalazinic, hypostictic and stictic 

acids. Predominant major compounds are constictic, hypostictic and stictic acid. In 

‘Thelotrema’ zebrinum substances of the protocetraric acid chemosydrome can co-occur. It is 

the most common chemosyndrome in the non-ocellularioid taxa (in Ocellularia s. lat. it is 

very rare), detected in 36 species of all genera except Fibrillithecis, Pseudoramonia and 

Reimnitzia: 

Chapsa lordhowensis, C. megalophthalma, C. megaphlyctidioides, C. minor, C. 

phlyctidioides, C. platycarpa, Chroodiscus australiensis, C. parvisporus, Leucodecton 

albidulum, L. compunctellum, L. glaucescens, L. subcompunctum, Melanotopelia rugosa, 

Myriotrema desquamans (strain I), M. eminens, M. phaeosporum, M. trypaneoides, 

Nadvornikia hawaiensis, Thelotrema alboolivaceum, T. bicinctulum, T. capetribulense, T. 

crassisporum, T. cupulare, T. cyphelloides, T. leucophthalmum, T. myriocarpum, T. 

porinoides, T. thesaurum, T. triseptatum, Topeliopsis azorica (strain I), T. darlingtonii, T. 

elixii, T. kantvilasii, T. tasmanica, ‘Thelotrema’ guadeloupensis (strain I), ‘Thelotrema’ 

zebrinum. 

Norstictic acid chemosyndrome 

Includes variable concentrations of connorstictic, norstictic and subnorstictic acids, rarely 

with co-occurring substances of the stictic acid chemosyndrome (α-acetylconstictic, 

hyposalazinic and stictic acid). The predominant major compound is norstictic acid. Detected 

in six species of the genera Leucodecton, Myriotrema and Thelotrema: 

Leucodecton occultum, Myriotrema frustillatum, Thelotrema bicavatum, T. eungellaense, 

T. gallowayanum, T. porinaceum.


Protocetraric acid chemosyndrome 

Includes variable concentrations of conprotocetraric, fumarprotocetraric, protocetraric, 

succinprotocetraric and virensic acids. Predominant major compounds are fumarprotocetraric 

and protocetraric acid. Detected in five species of Chapsa, Pseudoramonia and the 

‘Leptotrema’ schizoloma-group (predominantly found in the genus Ocellularia): 

Chapsa asteliae, C. lamellifera, Pseudoramonia richeae, Thelotrema sp. I, ‘Thelotrema’ 

zebrinum (see also under stictic acid chemosyndrome). 

Hypoprotocetraric acid chemosyndrome 

Detected in two Myriotrema species, M. polytretum and M. viridalbum. Includes variable 

concentrations of conhypoprotocetraric, conprotocetraric, convirensic, 4-0-demethylnotatic, 

2-hydroxyhypoprotocetraric* (major in M. polytretum), 2-hydroxynornotatic* and 

hypoprotocetraric (major in M. viridalbum) acids. This chemosyndrome is more common in 

Ocellularia s. lat. 

Psoromic acid chemosyndrome 

Includes variable concentrations of 2'-0-demethylpsoromic, psoromic and subpsoromic 

acids. The predominant major compound is psoromic acid. In some specimens of 

Fibrillithecis halei an (additional) unknown sterol occurs, the 'platysporum unknown'*. The 

psoromic acid chemosyndrome is more common in Ocellularia and Stegobolus, but was also 

detected in eight species of Fibrillithecis, Myriotrema and Thelotrema: 

Fibrillithecis halei, Myriotrema clandestinum, M. glaucophaenum, M. microporum, M. 

rugiferum, M. temperatum, Thelotrema foveolare, T. saxicola. 

Olivaceic acid chemosydrome 

In Myriotrema olivaceum, two chemical strains were detected, including variable 

concentrations of O-methylolivaceic*, norisonotatic, norsubnotatic and olivaceic* acids. 

Strain II (with O-methylolivaceic and norisonotatic acids) might be conspecific with M. 

subterebrans (Frisch 2006). Norisonotatic and norsubnotatic acid are also known for several 

taxa in Ocellularia s. lat. 

Xanthones 

Lichexanthone 

Lichexanthone is the only known xanthone occurring in Graphidaceae, here it was found in 

Myriotrema viridalbum and Thelotrema sp. III. Predominantly found in Melanotrema and 

Redingeria, but also known from other taxa of Myriotrema and Stegobolus. 

Anthraquinones 

Two unknown anthraquinones, both turning purple in KOH, are found in Chroodiscus 

australiensis and Leptotrema wightii, respectively. In the latter species, deep red 

anthraquinone crystals are found clustered in the medulla, in TLC detectable as a low, grayish 

spot (after charring) with Rf-values of 33/14/14 (solvent systems A/B'/C). In C. australiensis 

the pigment crystals are orange to rust-red and scattered on the disc and the ascomata margin. 

In TLC, no spots were seen, probably due to low concentrations. 

Unknown substance 

In a chemical strain of ‘Thelotrema’ guadeloupensis the ‘cinchonarum unknown’* 

compound was detected. This substance often co-occurs with hirtifructic and conhirtifructic 

acid (=’diacida unknowns’) and is known for several Ocellularia species, but was also found 

in Myriotrema.


Chromatographic behavior of new or unknown substances 

Table 1: TLC characteristics of the four hitherto unknown substances (according to Elix, pers. com.) 

and for the ‘cinchonarum and platysporum unknowns’. 

Rf values for 

solvent system: 

Compound name A B' C Spot characteristics (after charring) 

2-hydroxyhypoprotocetraric 

acid 

12 29 7 dark blue to dark gray or black 

2-hydroxynornotatic acid 7 25 4 dark blue to dark gray 

O-methylolivaceic acid 7 23 17 pale purplish to grayish-brown 

olivaceic acid 2 14 4 pale purplish to grayish-brown 

‘cinchonarum unknown’ 2 7 3 (dark) gray 

'platysporum unknown' 53 57 60 

(dark) brown with yellowish to 

greenish aurora under UV light 

The following previously informal compound names used by several authors (e.g. Frisch, 

2006; Hale, 1974a) could be assigned to the following secondary metabolites: 

‘neoterebrans unknown’ =O-methylolivaceic acid 

‘olivaceum unknowns’: 

-high =norisonotatic acid 

-medium =norsubnotatic acid 

-low =olivaceic acid 

2. 6. Ecology 

The ecology of thelotrematoid lichens is poorly known, and the following is mainly based 

on field observations and the available data from the collection labels. A more detailed 

summary of the general ecology and habitat preferences of thelotremataceaen Graphidacae 

can by found in Frisch (2006). General treatments of the diversity and ecology of the lichen 

biota in tropical rainforests are provided by Coppins & Wolseley (2002), Galloway (1991), 

Lakatos & al. (2006) and Sipman & Harris (1989). Information on the substrate ecology of 

lichens is given by Armostrong (1988) and Brodo (1973). 

2. 6. 1. Substrate 

The majority of taxa occur on bark of various trees and shrubs. However, several species 

and some genera have a preference for other substrates, such as wood, leaves, bryophytes, 

detritus, soil and siliceous rock. Members of the genus Chroodiscus are strictly foliicolous, 

Melanotopelia and many species of Topeliopsis s. str. are found on bryophytes, where they 

often extend onto adjacent substrates (bark, wood, rock). Several (sub)alpine Tasmanian taxa


show a preference for exceptional substrates and in two cases are substrate specific: Chapsa 

asteliae is only known from dead leafs of Astelia alpina, Pseudoramonia richeae grows 

exclusively on bark and dead leafs of Richea scoparia; some specimens of Topeliopsis 

tasmanica and Chapsa lamellifera were found inhabiting plant debris or peaty soil. The latter 

is further the only known species of Chapsa in Australia that is also lignicolous or 

muscicolous. Other facultative muscicolous species (which, however, initially settle on bark 

and subsequently overgrow adjacent bryophytes) include Fibrillithecis halei, Leptotrema 

wightii and Reimnitzia santensis. In Thelotrema a relatively high number of facultative 

saxicolous taxa are found (in Australia: T. conveniens, T. porinaceum, T. saxatile, T. 

saxicola). 

2. 6. 2. Habitat 

The highest species diversity of thelotrematoid lichens is found in tropical coastal, lowland 

to mid-range rainforests and subtropical rainforests in predominantly higher altitudes, where 

they mainly occur in the upper forest levels or on other more open localities with better light 

conditions as for example along creeks. At these sites, they often form the dominant elements 

in the epiphytic mycobiota, and sometimes extensive colonies are found. It could be observed 

that larger thalli were often formed by members of Myriotrema and some taxa of Thelotrema, 

whereas other species of the latter genus as well as Chapsa species generally are found in 

smaller individuals, and a noticeable number seem to preferable pioneer on younger trees, 

shrubs or on thinner twigs respectively. In those habitats, often a high density of various 

competitive species (mostly with Graphidaceae s. str.) is found. Secondary or disturbed 

forests show a significant decrease in diversity, as well as drier localities as monsoon forests 

or wet sclerophyll forests. Only in tropical mangroves, certain species (e.g. Fibrillithecis 

halei, Leucodecton occultum, L. subcompunctum, Myriotrema phaeosporum, M. rugiferum, 

Thelotrema bicinctulum, T. lacteum, T. pachysporum) are found, whereas subtropical 

mangroves lack any thelotrematoid species. Leptotrema wightii is the only species that also 

tolerates seasonally dry habitats, otherwise, no other member of thelotremataceaen 

Graphidaceae was found in the Pacific hinterland of Queensland. 

The rainforests of the southern Pacific regions are poorer in species, however, many 

temperate species are only found there (e.g. Chapsa megalophthalma, Thelotrema lepadinum, 

‘Thelotrema’ guadeloupense, Topeliopsis decorticans). In the (sub)alpine forests of 

continental Australia (southern Great Dividing Ranges) thelotrematoid lichens are absent 

probably due to the low temperatures in winter, whereas the oceanic influenced (sub)alpine 

reaches of Tasmania are relatively rich in species. As noted by Frisch (2006) one of the 

reasons for the poorness of species diversity in temperate zones could lie in the increased 

competition caused by bryophytes and macrolichens. This might also be an explanation for 

the accumulative occurrence of muscicolous taxa in temperate zones, particularly of the genus 

Topeliopsis. In Tasmania, various habitats harbor thelotrematoid lichens, amongst them a 

high number of endemic species. Besides in rainforests, also in heath- and moorlands 

primarily members of Chapsa, Thelotrema, Topeliopsis and the Leptotrema schizoloma-group 

were found. 

2. 7. Distribution 

As shown in figure 3, trentepohlioid thelotremataceaen Graphidaceae occur in coastal 

Northern Territory, Pacific Queensland and New South Wales, Norfolk Island, southern 

Victoria, south-west Western Australia and Tasmania. Roughly five major distribution centers 

can be distinguished: the wider Cairns area (from Cape Tribulation to Townsville) in tropical


northern Queensland and the Proserpine/Mackay area in tropical north-central Queensland; 

the northern Sunshine Coast area and the wider Brisbane area (northern Brisbane district to 

Gibraltar Ranges) in subtropical southern Queensland and northern New South Wales; and 

the western Tasmanian ranges (cool-temperate). (It should be mentioned that many of the 

collections from Brisbane date back to the late 19 th century.) Smaller distribution centers are 

found in the Iron Ranges National Park (Cape York Peninsula, Queensland), and in New 

South Wales the Coffs Harbour/Taree area, the Barrington Tops area and the Sydney area 

(mainly Blue Mountains and Royal National Park). 

Fig. 3. Combined Australian distribution map of all treated species. [The gray tones of the circles 

indicate the number of species found in the region according to the legend. Insert upper right: 

Norfolk Island, insert lower left: southwest Western Australia (the latter also applies for the 

distribution maps used in the species descriptions)]. 

For Australia, the 103 known taxa can be grouped in seven different distributional types 

(table 2): a) strictly tropical (the largest group with 34 species), b) tropical to subtropical (28 

species), c) tropical to warm-temperate (11 species), d) tropical to cool-temperate (1 species), 

e) strictly subtropical (8 species), f) subtropical to cool-temperate (12 species) and g) strictly 

cool-temperate (9 species).


Table 2: Australian distribution types of the treated species. (For an explanation of the code see text 

above; *: does also occur in the tropics at high altitudes.) 

a) tropical 

b) tropicalsubtropical 

c) tropicalwarm-temperate 

d) tropicalcool-temperate 

e) subtropical 

f) subtropicalcool-temperate 

g) cooltemperate 

Taxa 

Chapsa astroidea, C. halei , C. indica, C. lassae, C. leprieurii, C. megaphlyctidioides, 

C. niveocarpa, C. platycarpa, C. tibellii, Chroodiscus australiensis, C. parvisporus, 

Myriotrema desquamans, M. eminens, M. myrioporum, M. polytretum, M. protoalbum, 

M. subconforme, M. viridialbum, Reimnitzia santensis, Thelotrema adjectum, 

T. alboolivaceum, T. capetribulense, T. crassisporum, T. eungellaense, T. foveolore, 

T. lepadodes, T. monosporum, T. myriocarpum, T. polythecium, T. thesaurum, 

T. triseptatum, Topeliopsis laceratula, T. pseudoexanthismocarpa, “Thelotrema” 

parvizebrinum 

Chapsa alborosella, C. leprocarpa, C. phlyctidioides, C. pulchra, Fibrillithecis halei, 

Leucodecton albidulum, L. compunctellum, L. glaucescens, L. occultum, Myriotrema 

album, M. clandestinum, M. frustillatum, M. glaucophaenum, M. microporum, 

M. olivaceum, M. phaeosporum, M. rugiferum, M. trypaneoides, Nadvornikia 

hawaiensis, Thelotrema bicinctulum, T. cupulare, T. gallowayanum, T. leucophthalmum, 

T. oleosum, T. pachysporum, T. saxatile, T. saxicola, T. subadjectum 

Leptotrema wightii, Leucodecton subcompunctum, Thelotrema conveniens, T. defossum, 

T. diplotrema, T. lacteum, T. nureliyum, T. porinaceum, T. porinoides, T. pseudosubtile, 

T. rugatulum 

Thelotrema bicavatum 

Myriotrema temperatum, Thelotrema crespoae, T. circumscriptum, T. cyphelloides, 

T. nostalgicum, Topeliopsis darlingtonii, T. elixii, “Thelotrema” subzebrinum 

Chapsa lordhowensis, Thelotrema megalophthalma, T. lepadinum, T. subtile, 

T. suecicum*, Topeliopsis acutispora, T. azorica, T. decorticans, T. subdenticulata, 

“Thelotrema” guadeloupensis, “Leptotrema” schizoloma*. “Thelotrema” zebrinum* 

Chapsa asteliae, C. lamellifera, C. minor, C. subpatens, Melanotopelia rugosa*, 

Pseudoramia richeae, Topeliopsis kantvilasii, T. muscigena, T. tasmanica 

Regarding the worldwide distribution, 31 of the species are so far known to be endemic to 

Australia [A], 6 species are Australasian [B], 3 species subantarctic [C], 21 species 

paleotropical [D], two species paleosubtropical [E], 39 species pantropical [F], one species 

pantemperate [G], and two species subcosmopolitan. [H] (the bold letters indicate the code 

for table 3). 

Table 3: Distributional types of the treated species. (The suffix ‘+’ indicates an extension into subtropical 

zones, ‘++’ indicates an extension into temperate zones.) 

[A] endemic 

[B] Australasian 

Taxa 

Chapsa asteliae, C. halei, C. lassae, C. lordhowensis, C. megaphlyctidioides, 

C. minor, C. niveocarpa, C. pulchra, C. tibellii, Myriotrema frustillatum, 

M. temperatum. Pseudoramia richeae, Thelotrema crespoae, T. capetribulense, 

T. crassisporum, T. cyphelloides, T. eungellaense, T. gallowayanum, T. oleosum, 

T. pseudosubtile, T. subadjectum, T. thesaurum, T. triseptatum, Topeliopsis 

acutispora, T. darlingtonii, T. decorticans, T. elixii, T. kantvilasii, T. tasmanica, 

“Thelotrema” parvizebrinum, “Thelotrema” subzebrinum 

Thelotrema monosporum, Chapsa lamellifera, C. megalophthalma, Leucodecton 

albidulum, Thelotrema circumscriptum, “Thelotrema” zebrinum


Taxa 

[C] subantarctic Melanotopelia rugosa, Topeliopsis subdenticulata 

[C+] subantarctic “Leptotrema” schizoloma 

[D] paleotropical 

[D+] paleotropical 

Chapsa indica, Myriotrema desquamans, M. eminens, M. polytretum, M. protoalbum, 

M. subconforme, Thelotrema polythecium, Topeliopsis laceratula, 

T. pseudoexanthismocarpa 

Myriotrema microporum, M. phaeosporum, Thelotrema bicinctulum, T. cupulare, 

T. foveolore, T. nostalgicum 

[D++] paleotropical Chapsa subpatens, Thelotrema nureliyum, T. rugatulum, Topeliopsis muscigena 

[E++] 

paleosubtropical 

[F] pantropical 

[F+] pantropical 

[F++] pantropical 

[G] pantemperate Topeliopsis azorica 

Thelotrema bicavatum, Thelotrema porinaceum 

Chapsa astroidea, C. leprieurii, Chroodiscus australiensis, C. parvisporus, 

Myriotrema myrioporum, M. viridialbum, Reimnitzia santensis, Thelotrema adjectum, 

T. alboolivaceum, T. lepadodes, T. myriocarpum 

Chapsa alborosella, C. leprocarpa, C. phlyctidioides, C. platycarpa, Fibrillithecis 

halei, Leucodecton compunctellum, L. glaucescens, L. occultum, Myriotrema album, 

M. clandestinum, M. glaucophaenum, M. olivaceum, M. rugiferum, M. trypaneoides, 

Nadvornikia hawaiensis, Thelotrema leucophthalmum, T. pachysporum, T. saxatile, 

T. saxicola 

Leptotrema wightii, Leucodecton subcompunctum, Thelotrema conveniens, 

T. defossum, T. diplotrema, T. lacteum, T. porinoides, T. subtile, “Thelotrema” 

guadeloupensis 

[H] subcosmopolitan Thelotrema lepadinum, Thelotrema suecicum 

2. 8. Key to the treated species 

1a Growing on dead or living leaves (foliicolous) ............................................................ 2 

1b Growing on different substrate than leaves (corticolous, lignicolous, muscicolous, 

saxicolous, terricolous or humicolous) ......................................................................... 5 

2a Occurring in alpine habitats in Tasmania, growing on Astelia or Richea, ascospores 

longer than 12 µm, containing the protocetraric acid chemosydrome ........................... 3 

2b Tropical, ascospores up to 12 µm long, containing the stictic acid chemosydrome........ 4 

3a Ascomata Geaster-like, not stipitate .......................................................Chapsa asteliae 

3b Ascomata distinctly stipitate .....................................................Pseudoramonia richeae 

4a Discs reddish, ascospores up to 10 µm long, with 2(3) loci ... Chroodiscus australiensis 

4b Discs brownish, ascospores up to 12 µm long, with 2-4 loci .. Chroodiscus parvisporus 

5a Ascomata mazaedious, ascospores brown, bilocular ............... Nadvornikia hawaiensis 

5b Ascomata without mazaedium, ascospores with more than 2 loci, or if bilocular, 

unpigmented................................................................................................................. 6


6a Ascomata distinctly stipitate, endemic to (sub)alpine heathland shrubs in Tasmania ....... 

................................................................................................. Pseudoramonia richeae 

6b Ascomata immersed to strongly emergent but without stipes ........................................ 7 

7a Ascospores transversely septate, rarely with a single longitudinal septum .................... 8 

7b Ascospores (eu)muriform or submuriform ................................................................. 60 

8a Ascospores brown at maturity ...................................................................................... 9 

8b Ascospores hyaline throughout or yellowish to brownish only in over-mature or 

deceased ascospores .................................................................................................. 15 

9a Ascospores 1 per ascus ................................................................ Thelotrema crespoae 

9b Ascospores 4-8 per ascus ........................................................................................... 10 

10a Lateral paraphyses absent ........................................................ Leucodecton albidulum 

10b Lateral paraphyses present ......................................................................................... 11 

11a Ascospores with 4-8 loci ........................................................................................... 12 

11b Ascospores with 8-26 loci .......................................................................................... 14 

12a Pores small, up to 100(200) µm in diameter, proper exciple fused .................................. 

................................................................................................... Topeliopsis kantvilasii 

12b Pores wide to gaping, exceeding 200 µm in diameter, proper exciple fused to free .... 13 

13a Species lacking secondary compounds, proper exciple fused to indistinctly free ............. 

........................................................................................................... Chapsa leprieurii 

13b Species containing the stictic acid chemosydrome, proper exciple free ........................... 

......................................................................................................... Chapsa platycarpa 

14a Ascospores 30-75 µm long, with 8-18(20) loci, loci oblong ............................................ 

.............................................................................................. Thelotrema pachysporum 

14b Ascospores (50)70-110(130) µm long, with (12)16-24(26) loci, cell walls usually 

becoming distinctly crenate, loci globose ....................................... Thelotrema lacteum 

15a Proper exciple dark-brown to carbonized, thalline rim becoming distinctly layered, 

ascospores non-amyloid, 30-80 µm long ..................................... Thelotrema zebrinum 

15b Proper exciple different or if dark and/or thalline rim layered, ascospores with amyloid 

reaction and/or smaller .............................................................................................. 16 

16a Mature ascospores non-amyloid ................................................................................. 17 

16b Mature ascospores amyloid ........................................................................................ 28 

17a Ascospores up to 35 µm, with up to 11 loci ............................................................... 18 

17b Ascospores in average longer than 35 µm, with 10-35 loci ......................................... 26 

18a Pores small, never exceeding 120 µm in diameter, ascomata not chroodiscoid........... 19 

18b Pores wide to gaping or ascomata distinctly chroodiscoid .......................................... 20 

19a Lateral paraphyses and secondary compounds lacking ............ Myriotrema protoalbum 

19b Lateral paraphyses present, containing the stictic acid chemosydrome ............................ 

................................................................................................ Thelotrema bicinctulum


20a Cool-temperate species ........................................................................... Chapsa minor 

20b (Sub)tropical species ................................................................................................. 21 

21a Thallus ecorticate, surface dull and roughened ........................................................... 22 

21b Thallus corticate, surface shiny to waxy and smooth .................................................. 24 

22a Discs epruinose or indistinctly pruinose, ascospores not exceeding 13 µm, with 3-6 loci 

.................................................................................................................. Chapsa halei 

22b Discs distinctly pruinose, ascospores up to 25 µm, with 6-9 loci ................................ 23 

23a Older ascomata becoming distinctly chroodiscoid, ascospores halonate, cell walls thin, 

lacking secondary compounds ........................................................ Chapsa alborosella 

23b Older ascomata rather indistinctly chroodiscoid, ascospores non-halonate, cell walls 

moderately thick, containing the stictic acid chemosydrome ....... Chapsa phlyctidioides 

24a Ascomata inconspicuous, up to 600 µm in diameter, disc rather indistinctly pruinose 

and brownish, thallus margin epruinose, off-white to reddish-brown, ascospores not 

exceeding 15 µm and with up to 5 loci .................................................... Chapsa lassae 

24b Ascomata conspicuous, exceeding 600 µm in diameter, disc and inner thallus margin 

±distinctly pruinose and bright, ascospores exceeding 15 µm and with more than 5 loci . 

................................................................................................................................... 25 

25a Hymenium up to 80 µm high, lateral paraphyses inconspicuous, ascospores up to 20(23) 

µm long, with up to 7(8) loci, lacking secondary compounds............. Chapsa astroidea 

25b Hymenium up to 70 µm high, lateral paraphyses conspicuous, ascospores up to 18 µm 

long, with up to 6(8) loci, containing the stictic acid chemosyndrome ............................. 

........................................................................................... Chapsa megaphlyctidioides 

26a Ascomata perithecioid to apothecioid, pores up to 100 µm, containing the stictic acid 

chemosyndrome ...................................................................... Topeliopsis darlingtonii 

26b Ascomata becoming chroodiscoid when mature, lacking secondary compounds ........ 27 

27a Ascospores not exceeding 70 µm and with up to 24 loci, cell walls thin Chapsa pulchra 

27b Ascospores up to 110 µm, with up to 35 loci, cell walls thick .................. Chapsa indica 

28a Ascospores exceeding 90 µm and 20 loci .................................................................. 29 

28b Ascospores up to 90(110) µm with up to 20(22) loci .................................................. 34 

29a Proper exciple fused and usually not visible from the surface ..................................... 30 

29b Proper exciple distinctly free and visible from the outside ......................................... 32 

30a Thallus thick, distinctly corticate, thalline rim usually eroded in apical parts, 

corticolous, tropical species................................. Topeliopsis pseudoexanthismocarpa 

30b Thallus thin, covered by protocortex or indistinctly corticate, thalline rim entirely 

eroded with age, predominantly muscicolous, sub-tropical to cool-temperate species . 31 

31a Ascospores up to 130(150) µm with up to 32 loci, form bacillar, often distinctly bent, 

loci angular and pinched also in mature stages .......................... Topeliopsis acutispora 

31b Ascospores up to 100(110) µm with up to 24(25) loci, form fusiform and straight, loci 

in mature stages roundish ................................................... Topeliopsis subdenticulata


32a Ascospores not exceeding 150(170) µm and with up to 26(28) loci, containing the 

stictic acid chemosyndrome ....................................................... Thelotrema porinoides 

32b Ascospores up to 240 µm, with up to 38 loci, lacking secondary compounds or 

containing the protocetraric acid chemosyndrome....................................................... 33 

33a Ascospores 4-8 per ascus, lacking secondary compounds .......... Thelotrema nureliyum 

33b Ascospores 2-4 per ascus, containing the protocetraric acid chemosyndrome .................. 

................................................................................................ Thelotrema nostalgicum 

34a Ascospores small, up to 13 µm, with up to 2 loci ................... Myriotrema myrioporum 

34b Ascospores exceeding 13 µm and with more than 2 loci ............................................ 35 

35a Ascospores exceeding 50 µm ..................................................................................... 36 

35b Ascospores up to 50 µm ............................................................................................ 38 

36a Ascospores non-amyloid (to faintly amyloid), thalline rim distinctly split and layered, 

proper exciple fused to apically exposed, containing the stictic acid chemosyndrome ..... 

................................................................................................. Topeliopsis darlingtonii 

36b Ascospores amyloid, thalline rim entire to slightly split, proper exciple ±free, lacking 

secondary compounds ................................................................................................ 37 

37a Ascospores up to 60 µm, with up to 16 loci, predominantly with crenate surface............. 

............................................................................................. Thelotrema pseudosubtile 

37b Ascospores up to 90(110) µm, with up to 20(22) loci, with entire surface........................ 

.................................................................................................. Thelotrema diplotrema 

38a Lateral paraphyses lacking, ascomata usually ±small, immersed or indistinctly emergent 

................................................................................................................................... 39 

38b Lateral paraphyses present, ascomata large and distinctly emergent or chroodiscoid .. 46 

39a Proper exciple predominantly distinctly free, at least in mature ascomata. .................. 40 

39b Proper exciple predominantly fused ........................................................................... 42 

40a Thallus unfissured or slightly fissured, ascomata often becoming distinctly emergent, 

thalline rim typically strongly split to lacerate or eroded, ascospores up to 18(20) µm 

with up to 6(8) loci .......................................................... Myriotrema glaucophaenum 

40b Thallus becoming ±distinctly fissured to areolate, ascomata predominantly immersed, 

ascospores up to 16(18) µm with up to 4(5)loci ......................................................... 41 

41a Containing the ‘olivaceum unknown’ compound (strain I or II), thallus up to 600 µm 

high, proper exciple in section brown and apically darkened, ascospores up to 15 µm 

long ........................................................................................... Myriotrema olivaceum 

41b Containing the psoromic acid chemosyndrome, thallus up to 800 µm high, proper 

exciple in section grayish and apically not darkened, ascospores up to 16(18) µm long. .. 

.............................................................................................. Myriotrema microporum 

42a Thallus thin, up to 100 µm high, ascomata typically elongated and arranged in rows, 

ascospores non-amyloid to faintly amyloid ............................. Myriotrema protoalbum 

42b Thallus thick, exceeding 100 µm in height, ascomata different, ascospores distinctly to 

strongly amyloid ........................................................................................................ 43


43a Thallus usually distinctly fissured, surface verruculose to verrucose, subtropical............. 

............................................................................................... Myriotrema temperatum 

43b Thallus unfissured, surface continuous, tropical to more rarely extending in the subtropics 

...................................................................................................................... 44 

44a Thallus up to 300 µm high, ascospores up to 18(22) µm long with up to 4 loci, 

containing the hypoprotocetraric acid chemosyndrome ............. Myriotrema polytretum 

44b Thallus exceeding 300 µm in height, ascospores longer than 18(22) µm (up to 25[28] 

µm), containing the psoromic acid chemosyndrome or lacking secondary compounds 45 

45a Lacking secondary compounds, thallus with abundant calcium oxalate crystals, 

ascospores frequently with a single, longitudinal septum ................. Myriotrema album 

45b Containing the psoromic acid chemosyndrome, thallus without or with rather sparse 

calcium oxalate crystals, ascospores seldom with longitudinal septum............................. 

............................................................................................ Myriotrema clandestinum 

46a Ascomata chroodiscoid, at least in older stages, proper exciple fused or indistinctly 

(apically) free (inner thalline rim layers sometimes might be confused with proper 

exciple!) ..................................................................................................................... 47 

46b Ascomata not chroodiscoid, proper exciple distinctly free ......................................... 50 

47a Thallus thin, up to 150 µm high, with thin (up to 10 µm high), incontinuous cortex or 

ecorticate, ascospores up to 30 µm and with up to 10 loci........................................... 48 

47b Thallus thick, up to 400 µm high, distinctly corticate, ascospores up to 45 µm and with 

up to 12(14) loci...................................................................................... Chapsa tibellii 

48a (Sub)tropical species, ascospores up to 24 µm long, with up to 8(x2) loci........................ 

................................................................................................... Chapsa phlyctidioides 

48b Cool-temperate species, ascospores up to 30 µm long, with up to 10 loci ................... 49 

49a Ascospores with thin cell walls, non-amyloid to faintly amyloid, containing the stictic 

acid chemosyndrome .............................................................................. Chapsa minor 

49b Ascospores with thick cell walls, distinctly to strongly amyloid, lacking secondary 

compounds ...................................................................................... Chapsa subpatens 

50a In older ascomata thalline rim becoming layered, proper exciple dark brown to slightly 

carbonized........................................................................... Thelotrema parvizebrinum 

50b Ascomata and proper exciple different or if thalline rim layered, proper exciple not 

dark, if proper exciple dark brown or carbonized, then only apically and thalline rim not 

layered ....................................................................................................................... 51 

51a Ascomata immersed ................................................................................................... 52 

51b Ascomata (at least in older stages) distinctly emergent .............................................. 55 

52a Ascospores 15-20 µm long, with 4 loci .................................... Thelotrema triseptatum 


53a Ascospores with thick cell walls, up to 40(45) µm long, with up to 14(16) loci, 

containing norstictic acid ........................................................... Thelotrema bicavatum 

53b Ascospores with thin cell walls, up to 35(40) µm long, with up to 11(12) loci, 

containing the stictic acid chemosyndrome or lacking secondary compounds ............. 54


54a Thallus ecorticate, ascospores with distinct halo, lacking secondary compounds.............. 

.................................................................................................... Thelotrema defossum 

54b Thallus corticate (cortex sometimes incontinuous), ascospores with thin, indistinct halo, 

containing the stictic acid chemosyndrome ............................. Thelotrema bicinctulum 

55a Ascospores up to 23 µm long, with 4-6 loci ........................ Thelotrema alboolivaceum 


56a Ascospores either becoming brownish in over-mature stages or with distinctly thick cell 

walls from early stages on, lacking secondary compounds ........................................ 57 

56b Ascospores hyaline throughout and with unthickened cell walls in all developmental 

stages, containing depsidones .................................................................................... 58 

57a Ascospores with thin cell walls in young stages, strongly amyloid, becoming brownish 

in over-mature stages, loci large ...................................................... Thelotrema subtile 

57b Ascospores with thick cell walls from early stages on, faintly amyloid, strictly hyaline, 

loci small .................................................................................... Thelotrema suecicum 

58a Thalline rim in older ascomata becoming split to slightly lacerate or eroded and often 

slightly layered, thallus indistinctly fissured and distinctly verrucose, containing the 

stictic acid chemosyndrome ................................................ Thelotrema capetribulense 

58b Thalline rim entire to slightly split, unlayered, thallus distinctly fissured, continuous to 

rarely distinctly verrucose, containing salazinic or norstictic acid .............................. 59 

59a Pores small, up to 200 µm in diameter, discs indistinctly pruinose, epihymenium thin, 

containing salazinic acid ................................................... Thelotrema circumscriptum 

59b Pores wide, up to 300(500) µm in diameter, discs distinctly pruinose, epihymenium 

thick, containing norstictic acid .................................................. Thelotrema bicavatum 

60a Ascospores 1-4 per ascus, eumuriform ....................................................................... 61 

60b Ascospores 4-8 per ascus, eumuriform to submuriform .............................................. 85 

61a Ascomata large, distinctly chroodiscoid, at least in older stages ................................. 62 

61b Ascomata small to moderately large, perithecioid to apothecioid ............................... 64 

62a Ascospores amyloid, containing the stictic acid chemosyndrome . Chapsa lordhowensis 

62b Ascospores non-amyloid, lacking secondary compounds ........................................... 63 

63a Hymenium und lateral paraphyses interspersed with fine granules, ascospores up to 190 

µm long, hymenium up to 200 µm high .......................................... Chapsa niveocarpa 

63b Hymenium clear (lateral paraphyses interspersed), ascospores up to 130 µm long, 

hymenium up to 140 µm high .......................................................... Chapsa leprocarpa 

64a Hymenium inspersed ................................................................................................. 65 

64b Hymenium clear ........................................................................................................ 67 

65a Ascospores hyaline, containing norstictic acid......................... Thelotrema porinaceum 

65b Ascospores hyaline or ±brownish at later maturity, lacking secondary compounds or 

containing the psoromic acid chemosyndrome............................................................ 66


66a Ascospores distinctly brownish only in over-mature stages, lacking secondary 

compounds ................................................................................... Thelotrema saxicola 

66b Ascospores distinctly brownish at maturity, containing the psoromic acid 

chemosydrome ............................................................................. Thelotrema oleosum 

67a Proper exciple distinctly carbonized or dark brown .................................................... 68 

67b Proper exciple non-carbonized, if dark, then only apically ......................................... 69 

68a Ascomata sessile, distinctly emergent, urceolate to subglobose, thalline rim margin 

rugose to entire, ascospores up to 230 µm, containing the stictic acid chemosyndrome ... 

.................................................................................................... Melanotopelia rugosa 

68b Ascomata erumpent, ±emergent, hemispherical to rather indistinctly urceolate, thalline 

rim margin coarsely split to lacerate, ascospores up to 120(130) µm, lacking secondary 

compounds or containing salazinic acid ................................ Thelotrema schizolomum 

69a Lateral paraphyses lacking ......................................................................................... 70 

69b Lateral paraphyses present (in perithecioid ascomata often inconspicuous) ................ 72 

70a Ascospores becoming distinctly brown, up to 130 µm long Leucodecton compunctellum 

70b Ascospores hyaline, up to 200 µm long ......................................................................71 

71a Ascospores non-amyloid, containing norstictic acid ................ Myriotrema frustillatum 

71b Ascospores amyloid, containing the stictic acid chemosyndrome .. Myriotrema eminens 

72a Thalline rim predominantly and in large parts distinctly split to lacerate or eroded, 

±pruinose, proper exciple fused ................................................................................ 73 

72b Thalline rim entire to slightly split, proper exciple free, at least in apical parts ........... 77 

73a Ascospores non-amyloid, 2-4(8) per ascus, up to 100 µm .......... Topeliopsis tasmanica 

73b Ascospores amyloid, 1-2 per ascus, up to 210 µm ...................................................... 74 

74a In older ascomata pores gaping to almost appearing chroodiscoid, ascospores hyaline, 

producing ascoconidia in over-mature stages, containing hypostictic and hypoconstictic 

acids as major compounds .................................................................. Topeliopsis elixii 

74b Pores rather small, only gaping in deceased, sterile ascomata, ascospores without 

ascoconidia, or if bearing ascoconidia, ascospores yellowish to brownish at late 

maturity, lacking secondary compounds or containing stictic acid as major compound 75 

75a Predominantly muscicolous, rarely corticolous, ascomata distinctly emergent, usually 

with reddish-brown base, ascospores up to 210 x 55 µm, becoming yellowish to 

brownish at late maturity and producing acoconidia ................... Topeliopsis muscigena 

75b Muscicolous, saxicolous or corticolous, ascomata immersed to emergent, ascospores up 

to 160(170) x 45 µm, hyaline and not producing ascoconidia ..................................... 76 

76a Corticolous, thallus dark and with distinct, thick, yellowish cortex, ascospores with 

thick cell walls ............................................................................ Topeliopsis laceratula 

76b Corticolous, muscicolous or saxicolous, thallus pale, cortex indistinct or lacking, 

ascospores with thin cell walls ........................................................ Topeliopsis azorica 

77a Ascospores up to 100 µm, becoming distinctly brown ............................................... 78 

77b Ascospores exceeding 100 µm, hyaline to yellowish or distinctly brown ................... 79


78a Ascospores 2-8 per ascus, becoming brown in rather late maturity, distinctly amyloid, 

with thick cell walls and ±distinctly tapered ends, loci large, usually undivided in ends 

of ascospores .............................................................................. Thelotrema lepadodes 

78b Ascospores 1-4 per ascus, brown at early maturity, non-amyloid to faintly amyloid, with 

thin cell walls and indistinctly tapered ends, loci small, usually divided in ascospore 

ends ...................................................................................... Thelotrema monosporum 

79a Ascospores up to 350(400) µm long, with short, distinctly tapered ends ......................... 

............................................................................................ Thelotrema gallowayanum 

79b Ascospores up to 230(250) µm long, ends different ................................................... 80 

80a Ascospores 2-4 per ascus, containing norstictic acid .............. Thelotrema eungellaense 

80b Ascospores 1 to rarely 2 per ascus, containing norstictic or psoromic acid or lacking 

secondary compounds ................................................................................................ 81 

81a Asci with thick walls and lacking tholus, ascospores distinctly amyloid, ascospore walls 

and endospore remaining non-amyloid , containing the psoromic acid chemosyndrome .. 

..................................................................................................... Thelotrema foveolare 

81b Asci and ascospores different, containing norstictic acid or lacking secondary 

compounds ................................................................................................................. 82 

82a Ascospores hyaline to pale yellowish in over-mature stages ...................................... 83 

82b Ascospores becoming distinctly pigmented with maturity .......................................... 84 

83a Ascomata perithecioid, pores up to 80 µm in diameter, containing norstictic acid ........... 

................................................................................................ Thelotrema porinaceum 

83b Ascomata perithecioid to more often apothecioid, pores exceeding 80 µm in diameter, 

lacking secondary compounds .................................................. Thelotrema rugatulum 

84a Ascospores non-amyloid, brown at early stages ............................. Thelotrema saxatile 

84b Ascospores amyloid, brown at late maturity .............................. Thelotrema conveniens 

85a Hymenium inspersed ............................................................. Myriotrema trypaneoides 

85b Hymenium clear ........................................................................................................ 86 

86a Proper exciple distinctly carbonized to dark brown .................................................... 87 

86b Proper exciple non-carbonized or if dark, then only apically ...................................... 89 

87a Ascospores up to 35 µm ........................................................ Thelotrema subzebrinum 

87b Ascospores exceeding 35 µm, up to 130 µm .............................................................. 88 

88a Ascospores 1-4(6) per ascus, up to 120(130) µm, lacking secondary compounds or 

containing salazinic acid ....................................................... Thelotrema schizolomum 

88b Ascospores 4-8 per ascus, up to 60(70) µm, containing the stictic acid chemosyndrome 

or ‘cinchonarum unknown’ compounds .............................. Thelotrema guadeloupense 

89a Ascospores exceeding 80 µm ..................................................................................... 90 

89b Ascospores up to 80 µm ............................................................................................ 93


90a Proper exciple fused, not visible from the outside, thalline rim lacerate, distinctly lobed 

and ±layered .............................................................................. Topeliopsis tasmanica 

90b Proper exciple ±free and visible from the outside, thalline rim different ..................... 91 

91a Ascospores becoming brownish .................................................. Thelotrema lepadodes 

91b Ascospores remaining hyaline ....................................................................................92 

92a Ascospores with thin cell walls, containing the stictic acid chemosyndrome ................... 

.................................................................................................. Thelotrema thesaurum 

92b Ascospores with thick cell walls, lacking secondary compounds Thelotrema lepadinum 

93a Ascospores becoming distinctly brownish ................................................................. 94 

93b Ascospores hyaline to yellowish .............................................................................. 100 

94a Thallus shiny, distinctly corticate, pores tiny to small, up to 30 µm in diam., proper 

exciple fused ......................................................................... Myriotrema desquamans 

94b Thallus predominantly dull and ecorticate, or shiny and corticate, then ascomata 

different ..................................................................................................................... 95 

95a Ascomata becoming chroodiscoid, lacking secondary compounds Reimnitzia santensis 

95b Ascomata perithecioid to apothecioid, containing stictic or norstictic acid, or an 

unknown anthraquinone (then thallus with reddish, K+ purple crystals) ..................... 96 

96a Thallus distinctly epiphloedal, thick, bulging away from substrate and with ±distinct 

reticulate pattern, calcium oxalate crystals conspicuous, abundant and often arranged in 

columns ......................................................................................................................97 

96b Thallus different, proper exciple predominantly free .................................................. 99 

97a Thallus with conspicuous reddish, K+ purple crystals, containing an unknown 

anthraquinone .................................................................................. Leptotrema wightii 

97b Thallus without red crystals, containing the stictic acid chemosyndrome ................... 98 

98a Ascomata clustered, often forming stromata-like structures, pores wide, disc visible, 

proper exciple predominantly free ........................................ Leucodecton glaucescens 

98b Ascomata arrangement differently, pores small, disc non-visible, proper exciple 

predominantly fused ............................................................ Myriotrema phaeosporum 

99a Ascospores becoming brownish at rather early stages, up to 35(40) µm long, containing 

norstictic acid ............................................................................ Leucodecton occultum 

99b Ascospores becoming brownish at late stages, up to 45(50) µm long, containing the 

stictic acid chemosyndrome ........................................... Leucodecton subcompunctum 

100a Ascospores with more than 12 x 4 loci ..................................................................... 101 

100b Ascospores up to 11(12) x 4 loci .............................................................................. 108 

101a Proper exciple fused ................................................................................................. 102 

101b Proper exciple becoming ±free, at least in older ascomata ........................................ 105 

102a Ascomata apothecioid, lacking secondary compounds .............. Topeliopsis decorticans 

102b Ascomata becoming chroodiscoid, containing the stictic or protocetraric acid 

chemosyndrome ...................................................................................................... 103


103a Ascomata margins weakly layered, warm-temperate to subtropical species, containing 

the stictic acid chemosyndrome, ............................................. Chapsa megalophthalma 

103b Ascomata margins strongly layered, cool-temperate species, containing the protocetraric 

acid chemosyndrome, ............................................................................................... 104 

104a Epiphytic on Astelia, ascomata up to 1.5 mm in diameter, ascospores amyloid ............... 

.............................................................................................................. Chapsa asteliae 

104b Substrate different, ascomata exceeding 1.5 mm in diameter (up to 3 mm), ascospores 

non-amyloid ................................................................................... Chapsa lamellifera 

105a In older ascomata pores wide to gaping, up to 1.5 mm in diameter, containing the stictic 

acid chemosyndrome, ...................................................... Thelotrema leucophthalmum 

105b Pores small to wide, up to 500 µm in diameter, lacking secondary compounds ........ 106 

106a Ascospores up to 135 µm, ascomata becoming distinctly emergent, subtropical to cooltemperate 

species....................................................................... Thelotrema lepadinum 

106b Ascospores up to 80 µm, ascomata immersed to indistinctly emergent, predominantly 

tropical species, extending into the subtropics .......................................................... 107 

107a Thallus rather thin, ascomata immersed, proper exciple distinctly free and visible, 

thalline rim not layered, ascospores up to 35 µm, cell walls thin, but endospore 

distinctly thickened ................................................................ Thelotrema subadjectum 

107b Thallus thick, ascomata immersed to slightly emergent, proper exciple often only 

indistinctly free and only partly visible, thalline rim becoming ±layered, ascospores up 

to 80 µm, cell walls thick, endospore not thickened ..................... Thelotrema adjectum 

108a Ascospores non-amyloid, lateral paraphyses present (sometimes inconspicuous) ..... 109 

108b Ascospores faintly to distinctly amyloid, lateral paraphyses present or lacking ........ 113 

109a Proper exciple fused, lacking secondary compounds ............... Thelotrema polythecium 

109b Proper exciple ±free, containing the stictic acid chemosyndrome ............................. 110 

110a Thalline rim becoming distinctly split to lacerate and lobed, ascomata chroodiscoid ....... 

..................................................................................................... Thelotrema cupulare 

110b Thalline rim entire, slightly split or somewhat eroded, ascomata apothecioid ........... 111 

111a Thallus ecorticate, ascomata immersed ................................... Thelotrema cyphelloides 

111b Thallus corticate, ascomata immersed to slightly or distinctly emergent ................... 112 

112a Thallus thin, ascomata large, up to 600 µm in diameter, ascospores up to 20 µm and 7 x 

4 loci .................................................................................... Thelotrema crassisporum 

112b Thallus moderately thick, ascomata small, up to 350 µm in diameter, ascospores up 

to30(40) µm and 11(12) x 4 loci ........................................... Thelotrema myriocarpum 

113a Lateral paraphyses present, ascospores only faintly amyloid, containing the stictic acid 

chemosyndrome ....................................................................................................... 114 

113b Lateral paraphyses lacking (fibrillous apical proper exciple might be confused for 

lateral paraphyses!), ascospores distinctly amyloid, chemistry different ................... 115 

114a Thallus hypophloedal and ecorticate ....................................... Thelotrema cyphelloides 

114b Thallus moderately thick and corticate................................... Thelotrema myriocarpum


115a Thallus thin, ascomata immersed, lacking secondary compounds ................................... 

.............................................................................................. Myriotrema subconforme 

115b Thallus moderately thick to thick, ascomata immersed to distinctly emergent, with 

secondary metabolites .............................................................................................. 116 

116a Pores becoming wide to gaping, exceeding 300 µm in diameter (up to 600 µm), 

containing the hypoprotocetraric acid chemosyndrome ........... Myriotrema viridialbum 

116b Pores small, up to 100(200) µm in diameter, chemistry different ............................. 117 

117a Ascomata large, conspicuous, up to 1(1.5) mm in diameter, proper exciple thick, 

bearing apical fibrillae, containing the psoromic acid chemosyndrome and/or 

‘platyspora unknown’ compounds ................................................... Fibrillithecis halei 

117b Ascomata small, inconspicuous, up to 400 µm in diam., proper exciple different, 

containing the psoromic acid chemosyndrome ........................... Myriotrema rugiferum 

2. 9. Genera 

The following table provides an overview of the genus delimiting characters in the here 

treated group. Accepted genera in thelotremataceaen Graphidaceae that are treated elsewhere 

include: Acanthotrema, Ampliotrema, Diploschistes, Gyrotrema, Ingvariella, Melanotrema, 

Ocellularia, Redingeria and Stegobolus. 

Table 4: Main genus delimiting characters and other features of the treated groups. Explanation of the 

code: X =applicable in most cases, X =applicable in fewer cases, - =not applicable. [*: for further 

explanations see part 2. 5. 2.; a : aseptate in C. anomalus; b : not tested in most species; c : the type 

species L. zollingeri contains the hypoprotocetraric acid chemosyndrome; d : paraphyses tips and ascus 

structure could not be observed due to the mazaedium; e : isidia-like structures often present, probably 

representing immature ascomata (see part 2. 9. 9.); f : presence of lateral paraphyses uncertain (see part 

2. 9. 10.); g : a ‘key-hole appearance’ of the ascus tips was reported for some specimen (see part 2. 9. 

10.); h : ascoconidia.] 

Substrate 

Chapsa 

Chroodiscus 

Fibrillithecis 

Leptotrema 

bark X - X X X X X X X X X X X 

rock X - - - - - - - - - X X - 

leaves X X - - - - - - X - - - - 

bryophytes X - X - - X - - - X - X - 

wood X - - - - - - - - - - X - 

soil X - - - - - - - - - - - - 

peaty soil/plant debris X - - - - - - - - - - X - 

Leucodecton 

Melanotopelia 

Myriotrema 

Nadvornikia 

Pseudoramonia 

Reimnitzia 

Thelotrema 

Topeliopsis 

L. schizolomagroup


Thallus 

Chapsa 

Chroodiscus 


Leptotrema 

-predominantly 

hyposubstratic X - X - X X X X X X X X X 

-predominantly 

episubstratic X X X X X X X X X X X X X 

-cortex: 

distinctly corticate (true 

cortex present) X - X - - X X - X - X X - 

ecorticate, with protocortex X X - X X - X X X - X X X 

ecorticate, cortex structures 

absent X - - - X - - X - X X - - 

-oxalate crystals: 

conspicuous, large, 

±columnar - - - X X - X - - X X - - 

-vegetative propagules: 

isidia present - X X - - - X - - e X X - - 

soralia present - - - - X - - X - - - - - 

Ascomata -morphology*: 

Geaster-like X X - - - - - - - X - - - 

mazaedious - - - - - - - X - - - - - 

stipitate - - - - - - - - X - - - - 

myriotremoid s. str. - - - X X - X - - - - - - 

thelotremoid s. str. - - - - - - X - - - X - - 

perithecioid-thelotremoid - - - - - - X - - - X - - 

emergent-perithecioid - - X - - - X - - - - - - 

topeliopsioid - - - - - X - - - - - X - 

layered-carbonized - - - - - - - - - - - - X 

-proper exciple: 

evanescent to rather thin X X - - - - X - - - X - - 

rather thick to distinctly 

thick X X X X X X X X X X X X X 

prosoplectenchymatous (to 

fibrillous) - - X - - - X - - X - - - 

paraplectenchymatous X X - X X X - X X - X X X 

rather pallid X X X - - - - X - - X X - 

rather strongly pigmented - - - X X - X - X X X X X 

distinctly carbonized - - - - - X - - X - - - X 

-hymenium: 

inspersion present 

in parts ±distinctly branched 

- - - - - - X - - - X X - 

(except epithecium) 

paraphyses tips distinctly 

- - - X X - X - - X X - - 

thick and/or irregular X - X X X - X ? d 

paraphyses tips regular, 

unthickened or slightly 

- X X - - 

thickened X X - - X X X ? d X - X X X 

lateral paraphyses present X - - - - X - - X X f X X X 

Leucodecton 

Melanotopelia 

Myriotrema 

Nadvornikia 

Pseudoramonia 

Reimnitzia 

Thelotrema 

Topeliopsis 

L. schizolomagroup


Conidia 

Chemistry 

Distribution 

Chapsa 

-ascus: 

walls thickened, tholus 

absent - - - - X - X ? d 

Chroodiscus 


Leptotrema 

Leucodecton 

Melanotopelia 

Myriotrema 

Nadvornikia 

Pseudoramonia 

Reimnitzia 

Thelotrema 

Topeliopsis 

L. schizolomagroup 

- - g X - X 

walls unthickened, tholus 

present (at least in younger 

stages) X X X - X X X ? d walls unthickened, tholus 

X X X X X 

absent X - - X - - - ? d 

ocular chamber conspicuous, 

- - - - - 

large, distinctly tapered X - - - - - X ? d 

-ascospores: 

- - - - - 

rather small (≤30 µm long) X X X X X - X X X X X X X 

very large (>130 µm long) X - - - - X X - - - X X - 

transseptate X X a 

- - X - X X X - X X X 

submuriform to eumuriform 

cell walls (and endospore) 

X X X X X X X - - X X X X 

rather thin 

cell walls (and endospore) 

X X - X - - X - X - X X X 

rather thick X - X - X X X X - X X X X 

hyaline X X X - X X X - X - X X X 

brown X - - X X X X X - X X X X 

non-amyloid X X - X X X X X X X X X X 

faintly to strongly amyloid X - X X X X X - - X X X X 

bacilliform - - X X X - X - - X - X h 

irregular, oblong or 

ellipsoid X - - - X - - - - - X X h 

fusiform - - - - - - X - - X - - - 

β-orcinol depsidones X X b X X c X X X X X - X X X 

xanthones - - - - - - X - - - X - - 

anthraquinones - X - X - - - - - - - - - 

tropical to subtropical X X X X X X X X - X X X X 

warm- to cool-temperate X - - X X X - - X - X X X 

2. 9. 1. Chapsa Massal., Atti Reale Ist. Veneto Sci. Lett. Arti, ser. 3, 5: 256 (1860). Type 

species: Chapsa indica Massal. 

Asteristion Leight., Trans. Linn. Soc. London 27: 163 (1869). Asteristium Clements, The genera of fungi: 76 

(1909). Thelotrema sect. Asteristion (Leight.) Matsum., J. Hattori Bot. Lab. 88: 16 (2000). Type species: 

Asteristion erumpens Leight. =Chapsa platycarpa (Tuck.) Frisch. 

- 

-


THALLUS – Crustose, very rarely partly bulging and flaking away from substrate, 

predominantly corticolous, rarely saxicolous, lignicolous, terricolous, humicolous, 

muscicolous or foliicolous. Predominantly ±thin, mainly hyposubstratic with minor 

episubstratic parts, c. 50-200 µm high, more rarely moderately thick and with ±distinct 

episubstratic parts, up to c. 400 µm high. Mostly pale, in shades of gray with greenish, 

tannish, yellowish or whitish tones, more rarely dark and in shades of olive with brownish or 

yellowish tones. Surface dull to distinctly shiny, rarely waxy, smooth to rough or pruinose, in 

predominantly hypophloedal thalli often with protuberant substrate structures, predominantly 

continuous to slightly verrucose or verruculose, rarely distinctly verrucose to verruculose. 

Predominantly unfissured, rarely distinctly fissured. Prothallus thin to indistinct, brown. 

Corticate to ecorticate, either without true cortex and covered by a c. 10-20 µm thick, 

discontinuous to continuous protocortex, rarely with distinctly conglutinated parts forming a 

true cortex; or covered by a c. 20-50 µm thick, hyaline to ±yellowish true cortex consisting of 

irregular to periclinal hyphae. Algal layer continuous to more rarely discontinuous, poorly to 

well developed, calcium oxalate crystals absent to abundant, small to large, scattered to 

clustered, rarely forming layers in lower thallus. Distinct medulla layer absent to more rarely 

present in thicker thalli and in ascomata area. Vegetative propagules not seen. 

ASCOMATA – Predominantly ±conspicuous, rarely inconspicuous (C. halei, C. subpatens), 

(moderately) large to very large, c. 0.6-5 mm in diam., roundish to ±irregular, particularly 

fused ascomata usually appearing distinctly irregular and/or ±branched, rarely somewhat 

elongated. Apothecioid in younger stages, becoming ±distinctly chroodiscoid with age, rarely 

indistinctly chroodiscoid throughout development. Erumpent, in regenerating ascomata often 

becoming ±sessile with successive ascomata generations. Mostly solitary to ±distinctly fused, 

rarely only marginally fused or strictly solitary, in some taxa ascomata ±frequently distinctly 

clustered in groups of several ascomata, then the individual ascomata often distinctly smaller 

than in solitary growing ascomata. In majority of species ascomata regenerating, then with 

±distinctly layered thalline rim. Immersed to slightly raised or with an annular margin, rarely 

distinctly emergent in regenerating ascomata, then depressed-urceolate to cupular. Disc 

entirely to partly visible from surface, at least in older stages, white, gray, flesh-colored or 

brownish, usually ±distinctly pruinose, more rarely epruinose. Proper exciple usually not 

visible from the surface to more rarely visible when becoming partly detached or in ascomata 

with strongly recurved thalline rims, ±whitish, rarely brownish, very rarely proper exciple 

distinctly visible from surface and forming an inner margin/pore. Thalline rim margin porelike 

only in younger stages or in ascomata with distinctly layered thalline rim, in older stages 

predominantly gaping, distinctly split and lobed to lacerate or eroded. Thalline rim unlayered 

to distinctly layered, epruinose to ±distinctly pruinose, whitish or brighter than thallus to more 

rarely brownish, yellowish or reddish-brown (usually due to protuberant substrate) internally 

and concolorous with thallus marginally, rarely concolorous with thallus throughout, 

predominantly erect to recurved, in layered margins often exfoliating. Proper exciple fused to 

slightly detached or apically exposed, very rarely distinctly free. Predominantly evanescent to 

thin, rarely distinctly thickened. Hyaline to rarely pale yellowish internally, yellow to brown, 

rarely orange marginally, in distinctly hypophloedal taxa often with substrate particles 

incorporated, apically sometimes more darkish and often covered by granules. Exciple nonamyloid 

to more rarely ±distinctly amyloid at the base. Subhymenium indistinct, evanescent 

to thin and with same color as basal exciple, rarely darkish brown to slightly carbonized, in 

some species with a distinct, hyaline, amyloid, lateral area. Hymenium non-amyloid, discoid 

to somewhat cupular, often with a ±broader base, up to c. 70-200 µm high, non-inspersed, 

predominantly clear, in C. niveocarpa distinctly interspersed, weakly to moderately, more 

rarely strongly conglutinated. Paraphyses un-thickened to rarely ±thickened, straight to 

slightly bent, parallel to slightly interwoven, unbranched, tips unthickened to ±distinctly 

thickened, sometimes ±distinctly irregular. Lateral paraphyses present, inconspicuous to


conspicuous, usually clearly separated from proper exciple, rarely not clearly separated 

(Topeliopsis-type), up to c. 20-45 µm long, predominantly clear, but sometimes also 

interspersed. True columella or columellar structures absent. Epihymenium (moderately) thin 

to (very) thick, predominantly hyaline, rarely pale yellowish or ±brownish, with fine to coarse 

granules. 

Asci 1-8-spored, non-amyloid, clavate, ascus walls unthickened (fide Frisch 2006 

distinctly thickened ascus walls in C. eitenii and C. zahlbruckneri), tholus predominantly 

present, in C. lamellifera absent, in younger stages predominantly thin to thick, becoming thin 

or not visible at maturity, rarely remaining distinctly thickened throughout development. 

Ascospores uni- to quadriseriate, very small to very large, 9-190 x 2-50 µm, predominantly 

transversely septate to more rarely (eu)muriform. Cell walls thin to moderately thick, more 

rarely distinctly thick, smooth to rarely crenate, in muriform ascospores endospore thin to 

moderately thick, non-halonate to distinctly halonate, especially in younger stages, thin to 

very thick, sometimes ±distinctly irregular. Ascospores hyaline to sometimes slightly 

yellowish or grayish at late maturity, rarely distinctly brown, non-amyloid to strongly 

amyloid. Oblong to ellipsoid or fusi- to claviform, rarely bacillar to bacillar-fusiform with 

roundish to acute ends, straight to more rarely ±bent, with 3-35 loci in transversely septate 

ascospores, with 8-18 x 1-6 or multiple loci in muriform ascospores, solitary end cells 

hemispherical to conical, loci small to large, roundish to angular, ±irregular, subglobose, 

oblong or lentiform, more rarely ± cuboid, transverse septae thin to ±distinctly thickened, 

regular to irregular, in densely muriform ascospores often distinct only younger stages, 

becoming indistinct and vanishing with age. 

PYCNIDIA – Only found in C. lordhowensis, see there for description. 

CHEMISTRY – Secondary compounds present or absent, if present, then predominantly of 

the stictic acid, rarely the protocetraric acid chemosydrome. 

ECOLOGY AND DISTRIBUTION – The Chapsa species in Australia predominantly occur on 

tree bark, rarely on dead wood, mosses, soil, siliceous rock, debris or on leaves in altitudes 

ranging between sea level and 1250 m. The greatest diversity of species is found in 

rainforests, rarely wet sclerophyll forests and monsoon forests, in tropical to sub-tropical 

climates of north-western Northern Territory, Pacific Queensland and northern New South 

Wales and on Lord Howe Island. Fewer species were found in rainforests, rarely wet 

sclerophyll forests and (sub)alpine heathlands, in warm- to cool-temperate climates in Pacific 

southern New South Wales, in southern Victoria and on Tasmania. At present state of 

knowledge, amongst the 19 species known in Australia, seven are endemic (C. halei, C. 

lassae, C. lordhowensis, C. megaphlyctidioides, C. niveocarpa, C. pulchra, C. tibellii), four 

are subantarctic (C. asteliae, C. lamellifera, C. megalophthalma, C. minor), two are 

paleotropical to paleotemperate (C. indica, C. subpatens) and six are pansubtropical to 

pantropical (C. alborosella, C. astroidea, C. leprieurii, C. leprocarpa, C. phlyctidioides, C. 

platycarpa). 

NOTES – This genus was recently resurrected (Frisch, 2006) to accommodate taxa formerly 

grouped in Chroodiscus and Thelotrema, here in particular the members of the ‘Thelotrema 

platycarpum-group’ (Salisbury, 1972b) and Thelotrema subgen. Asteristion (Matsumoto, 

2000). The genus name was introduced by Massalongo in 1860 and was neglected ever since. 

The taxa in Chapsa are characterized by a predominantly corticolous, thin thallus, typically 

large, chroodiscoid ascomata with fused to indistinctly free proper exciple (except C. 

platycarpa, which has a distinctly free proper exciple), un-branched, discoid to rarely slightly 

cupular hymenia and the presence of lateral paraphyses. The most similar genera are


Acanthotrema (not treated here, for differences see Frisch, 2006), Chroodiscus, Reimnitzia, 

Thelotrema and Topeliopsis. Chroodiscus is readily distinguished by smaller thalli, smaller 

ascomata without lateral paraphyses and a strictly foliicolous growth, Reimnitzia is 

distinguished by a thick, Leptotrema wightii-like, isidiate thalli with columnar calcium 

oxalate crystals, a slightly branched hymenium and probably absent lateral paraphyses (see 

observations there). Thelotrema and Topeliopsis predominantly differ by never distinctly 

chroodiscoid ascomata, although the distinction to Chapsa is sometimes difficult since certain 

taxa (e.g. T. cupulare, T. leucophthalmum, T. polythecium) also often have ±gaping apothecia 

with somewhat recurved margins. Most species in Thelotrema can be further distinguished by 

the distinctly free proper exciple and rather sessile than erumpent ascomata. The latter also 

applies for Topeliopsis, which further differs by a predominantly muscicolous growth, never 

distinctly discoid hymenia and usually thicker exciples with indistinctly separated lateral 

paraphyses. 

In the original resurrection (Frisch, 2006) the genus is further characterized by distinct and 

rigid paraphyses that have moniliform, branched or rarely simple tips and ascospores “of the 

thick-walled type” (ibid.). 

Given the generally great inter- and intraspecific variability in thelotremoid lichens, 

Chapsa can be considered a rather well separated group, which is also confirmed by the 

results of the molecular analyses (see part 3). Amongst the Australian taxa only the position 

of C. platycarpa is uncertain for the above mentioned morphological differences, however, in 

the single-gene molecular phylogeny presented by Frisch & al. (2006) it is well supported 

within Chapsa as a sister to C. indica. 

Ocellularia punicea (Müll.Arg.) Mangold & Lumbsch comb. nov. ined., known from 

southern Queensland was recently combined to Chapsa (Caceres, 2007). This taxon has 

ascomata with absent lateral paraphyses and a distinctly carbonized proper exciple and is thus 

rejected from the genus. 

Species descriptions: 

Chapsa alborosella (Nyl.) A. Frisch 

Bibl. Lichenol. 92: 91 (2006). Bas.: Graphis alborosella Nyl., Ann. Sci. Nat., Bot., sér. 4, 19: 372 (1863). 

Thelotrema alborosellum (Nyl.) Tuck., Gen. Lich.: 139 (1872). Ocellularia alborosella (Nyl.) Santesson, Symb. 

Bot. Upsal. 12(1): 308 1952). Chroodiscus alborosellus (Nyl.) Kalb, Lich. Neotrop. Fasc. XX (No. 456-475): 8 

(1991). Type: Colombia [Nova Granata], 2000 m, 1860, Lindig 2694 (H-Nyl.7635!-holotype; BM!-, FH-Tuck.!isotypes). 

ILLUSTRATION – Fig. 4. 

Thallus predominantly hypophloedal to slightly epiphloedal, very thin, up to c. 50 µm 

high, pale gray to grayish-green to pale yellowish-green. Surface dull to slightly shiny, 

smooth to slightly pruinose, often with protuberant substrate structures, continuous, 

unfissured. Cortex structures absent or covered by an incontinuous, very thin protocortex up 

to 10 µm thick. Algal layer continuous and well developed, calcium oxalate crystals sparse 

and scattered. Vegetative propagules not seen. Ascomata conspicuous, (moderately) large, up 

to c. 1.2 mm in diam., roundish to irregular, apothecioid when young, becoming chroodiscoid 

at maturity, erumpent, solitary to marginally fused, often clustered in groups of two to several 

ascomata, sometimes regenerating, immersed. Disc entirely to partly visible from surface, 

pale brown to grayish, distinctly pruinose. Proper exciple not visible from surface, thalline 

rim split, lobed to eroded, sometimes slightly layered, inside often pruinose, whitish or with


protuberant substrate, concolorous with thallus outside, slightly incurved to erect or recurved. 

Exciple fused, thin to evanescent, colorless internally to pale yellowish-brown or (pale) 

brownish marginally, apically often covered with fine grayish granules, non-amyloid. 

Hymenium up to c. 90 µm high, non-inspersed, strongly conglutinated, paraphyses ±straight, 

parallel to slightly interwoven, unbranched, slightly thickened apically, lateral paraphyses 

present, inconspicuous, up to c. 20 µm long. Epihymenium moderately thick, hyaline to pale 

grayish-brown, with grayish granules and crystals. Asci 8-spored, tholus thin, absent at 

maturity. Ascospores (very) small, transversely septate, cell walls (moderately) thin, 

±distinctly halonate, hyaline, non-amyloid, rarely oblong to usually fusi- to claviform with 

roundish to (sub-)acute ends, loci predominantly angular, end cells hemispherical to conical, 

septae thin, regular to irregular, 10-25 x 3-6 µm with 6-9 loci. Pycnidia not seen. 

Fig. 4. Chapsa alborosella: growth habit (A), ascomata (B) and ascospores (C, D). A.: Mangold 19 

zh; B., D.: FH-isotype; C.: Hale 830729. Bar= A: 1.2 mm; B: 0.8 mm; C: 10 µm; D: 15 µm. 

CHEMISTRY – Thallus K-, C-, PD-; no compounds detectable by TLC. 

ECOLOGY AND DISTRIBUTION – Chapsa alborosella was collected in Australia on tree bark 

in warm-temperate to tropical rainforests at an altitude of ca. 800 m. It is very rare in 

Australia, only known from two locations in Queensland and New South Wales. This is the 

first report for Australia, thus far it was known from the Neotropics (Hale, 1978, 1981;


Sipman, 1992), Africa (Frisch, 2006), India 

(Nagarkar & al., 1988), Sri Lanka (Hale, 1981) and 

Japan (Matsumoto, 2000), indicating that the 

species has a pantropical distribution and extends 

into subtropical regions. 

NOTES – This taxon is characterized by a thin, 

mostly hypophloedal, ecorticate thallus, pruinose 

apothecial discs and small, transversely septate, 

hyaline, non-amyloid ascospores with thin cell 

walls and the lack of secondary metabolites. 

Similar Australian species include C. astroidea and 

C. lassae, which, however, are readily distinguished 

by a thicker, corticate thallus and ±smaller 

ascospores. Chapsa halei also differs in having 

smaller ascospores (see also under this species). 

Another similar species is C. diploschistoides from 

Africa that differs in having a thicker, verrucose 

thallus less recurved ascomata margins and slightly 

larger ascospores (see also Frisch, 2006: 99 for a 

more detailed discussion). 

Thelotrema platycarpellum Vain. has been considered a synonym of C. alborosella by 

Hale (1981) and a distinct species by Frisch (2006), is regarded synonymous with C. 

astroidea here (see also under this species). 

SPECIMENS EXAMINED – Australia, Queensland, 18 km S of Ravenshoe on Tully Falls Rd., Hale 830729 

(US). New South Wales, Mt. Warning NP., Mangold 19ze, 19zh (F). 

Chapsa asteliae (Kantvilas & Vezda) Mangold comb. nov. ined. 

Bas.: Chroodiscus asteliae Kantv. & Vezda, Lichenologist 32: 328 (2000). Type: Australia, Tasmania, Mt. 

Curly, Kantvilas & Jarman 38/85 (BM!-isotype). 


Fig. 5. Australian distribution of 

C. alborosella. 

Thallus foliicolous, epi- to hyposubstratic, thin to very thin, up to c. 200 µm high, pale 

gray to pale tannish-gray. Surface slightly shiny, smooth, continuous to slightly verrucose, 

unfissured. True cortex present, ±continuous, up to c. 30 µm thick, consisting of periclinal 

hyphae. Algal layer ±continuous, weakly developed, calcium oxalate crystals absent. 

Vegetative propagules not seen. Ascomata conspicuous, large, up to c. 1.5 mm in diam., 

roundish, apothecioid to indistinctly chroodiscoid in older stages, sessile, solitary to 

marginally fused, regenerating, becoming distinctly emergent, flattened-subglobose to 

(flattened-)urceolate. Disc usually partly visible from surface, pale brownish, becoming darkgray 

with age, epruinose. Proper exciple usually not visible from surface, sometimes apically 

becoming somewhat visible as brownish line, thalline rim margin small to gaping, irregular to 

star-shaped, thalline rim becoming distinctly layered, radially split and ±lobed, concolorous 

with thallus or slightly darker, internally incurved to rarely slightly erect, outer layers 

becoming erect to more rarely recurved. Proper exciple predominantly fused, rarely apically 

exposed, thin, hyaline internally to pale brownish or grayish marginally, sometimes darkbrown 

apically, non-amyloid. Hymenium up to c. 150 µm high, non-inspersed, moderately 

conglutinated, paraphyses moderately thick, ±parallel, unbranched, tips moderately thick,


lateral paraphyses inconspicuous, up to 30 µm long, hypothecium conspicuous and thick, 

hyaline, strongly gelatinous and slightly amyloid. Epihymenium thin, hyaline, in older 

ascomata becoming distinctly brownish, rarely with few grayish granules. Asci 4-6(8)-spored, 

tholus thin, not visible at maturity. Ascospores moderately small to moderately large, (sub- 

)muriform, cell walls and endospore moderately thick, with thin to moderately thick halo, 

hyaline, weakly amyloid, oblong to roundish-fusiform with roundish to narrowed-roundish 

ends, loci large, predominantly irregular-roundish to slightly angular, subglobose to often 

elongate, transverse septae distinct and thick, ±regular, 30-60 x 10-15 µm with 8-16 x 1-4 

loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellowish-brown, C-, PD+ orange-red; containing succinprotocetraric 

(major), protocetraric and fumarprotocetraric acids (traces). 

Fig. 6. Chapsa asteliae: ascomata (A) and ascospores (B, C). A.-C.: BM-isotype. Bar= A: 1.2 mm; B: 

20 µm; C: 10 µm. 

ECOLOGY AND DISTRIBUTION – Chapsa asteliae 

grows on dead leaves of Astelia alpina in (sub-) 

alpine heathlands in altitudes ranging from 900 to 

1080 m. It is common in Tasmania and was 

previously only known from there. 


corticate bright thallus and flattened, subglobose to 

urceolate ascomata with (at least in older ascomata) 

distinctly layered margins and an often dark disc, 

moderately large, hyaline, muriform, amyloid, 

thick-walled and halonate ascospores with only a 

few longitudinal septae. Further it contains the 

protocetraric acid chemosyndrome. Chapsa 

lamellifera is similar, for differences see under this 

taxon. Chapsa asteliae has similarities to 

Topeliopsis, it differs from that genus, however, in 

having a thin exciple and thickened paraphyses tips. 

SPECIMENS EXAMINED – Australia, Tasmania, Hartz Mnt. 

NP., nr. Lady's Tarn, Mayrhofer 9643 (GZU). 


C. asteliae.


Chapsa astroidea (Berk. & Broome) Caceres & Lücking 

In Caceres, Libri Botanici 22: 51 (2007). Bas.: Platygrapha astroidea Berk. & Broome, J. Linn. Soc. (Bot.) 

14: 109 (1875). Ocellularia astroidea (Berk. & Broome) Hale, Mycotaxon 7: 377 (1978). Thelotrema astoideum 

(Berk. & Broome) Hale, Mycotaxon 11: 131 (1980). Type: Sri Lanka, Thwaites 629 (K!-lectotype, selected by 

Hale [1978: 377]). 

Ocellularia alba (Fée) Müll.Arg. var. caesiascens Räs., Arch. Soc. Zool. Bot. Fenn. Vanamo 3: 185 (1949). 

Type: Australia, North Queensland, Korunda, Aug.1893, Wilson s.n. (H-Räs.!-holotype, NSW!-isotype). 

Thelotrema platycarpellum Vain., Proc. Amer. Acad. Arts Sci. . 58: 138 (1923). Ocellularia platycarpella 

(Vain.) Zahlbr., Cat. Lich. Univ. II: 598 (1923). Chapsa platycarpella (Vain.) Frisch, Bibl. Lichenol. 92: 118 

(2006). Type: Trinidad & Tobago – Trinidad, Arima, Verdant Vale, Thaxter 57 (TUR-Vain. 26791-holotype). 


Fig. 8. Chapsa astroidea: ascomata (A, B) and ascospores (C, D). A., C.: H-holotype of O. alba var. 

caesiascens; B., D.: Hale 830623. Bar= A, B: 1 mm; C: 10 µm; D: 5 µm. 

Thallus epi- to hypophloedal, thin, up to c. 200 µm high, dark olive or olive-brown to pale 

yellowish-brown. Surface ±waxy, smooth, continuous, unfissured. True cortex present, 

±incontinuous, up to c. 30 µm thick, pale yellowish, consisting of periclinal to irregular 

hyphae. Algal layer continuous to incontinuous and poorly developed, calcium oxalate 

crystals abundant, predominantly clustered, sometimes forming layers within the medulla and 

the substratum. Vegetative propagules not seen. Ascomata conspicuous, large, up to c. 2.3 

mm in diam., roundish to slightly irregular to distinctly irregular in fused ascomata, distinctly 

chroodiscoid, erumpent, solitary to fused, sometimes clustered in small groups, sometimes 

regenerating, immersed. Disc usually entirely visible from surface, grayish, distinctly 

pruinose, often glittering. Proper exciple not visible from surface, thalline rim split, distinctly 

lobed to more rarely slightly eroded, rarely layered, internally pruinose, whitish, concolorous 

with thallus outside, erect to recurved. Exciple fused, thin to evanescent, colorless internally 

to pale yellowish-brown marginally, apically often covered with grayish granules, nonamyloid. 

Hymenium up to c. 80 µm high, non-inspersed, strongly conglutinated, paraphyses


±straight, parallel to slightly interwoven, unbranched, tips not thickened to slightly thickened, 

lateral paraphyses present, often inconspicuous, up to c. 30 µm long, subhymenium often 

conspicuously dark-brown to slightly carbonized. Epihymenium (moderately) thick, hyaline, 

with grayish granules and often ±large calcium oxalate crystals. Asci 8-spored, tholus thin, 

not visible at maturity. Ascospores (very) small, transversely septate, cell walls (moderately) 

thin, ±distinctly halonate, hyaline, non-amyloid, fusi- to claviform, often conspicuously 

narrow, with (sub-)acute ends, loci predominantly angular, often ±longitudinal elongated, end 

cells conical, septae thin, regular, 9-20(23) x 3-5µm, with 3-7(8) loci. Pycnidia not seen. 


E COLOGY AND D ISTRIBUTION – Chapsa 

astroidea was collected in Australia on tree bark in 

tropical rainforests in altitudes ranging from 30 to 

900 m. It is regionally common in Australia, 

occurring in northern Queensland. Outside of 

Australia it is known from Trinidad-Tobago, Africa 

(as C. platycarpella [Frisch, 2006]) and Sri Lanka 

(Hale, 1978, 1981) indicating a pantropical 

distribution. 

N OTES – This taxon is characterized by a 

corticate, dark thallus that gives a conspicuous 

contrast to the bright ascomata. Further it has a 

characteristic, usually dark subhymenial layer, 

small, narrow, transversely septate, hyaline, nonamyloid 

ascospores with thin cell walls. Chapsa 

lassae is a similar corticate species in Australia, it 

can be readily distinguished by the less distinctly 

pruinose, brownish discs, the off-white to reddishbrown, 

epruinose thalline margins and smaller 


C. astroidea. 

ascospores (up to 15 µm, with up to 5 loci). Other similar Australian species include C. 

alborosella and C. halei, both differing by a dull, ecorticate thallus and C . 

megaphlyctidioides, which contains stictic acid. Thelotrema platycarpella is included as a 

synonym although the type material could not be studied, since the detailed description and 

illustrations provided by Frisch (2006) agree with the material studied here. 

SPECIMENS EXAMINED – Australia, Queensland: Cape Tribulation, 3 km NE of Daintree River Crossing, Hale 

831668 (US). Atherton Tablelands: Junrum Environmental Park, nr. Kuranda, Hale 831433 (US); Palmerston 

NP., 7 km E of the west boundary, Hale 831435 (US); Mt. Hypipamee NP., S of Atherton, Hale 831922 (US); 

Ellingea Falls, 10 km E of Millaa Millaa, Lumbsch & Mangold 19144 b (F); Off Palmerston Hwy, 11 km from 

main hwy and c. 2 km N of S Johnstons Forestry Camp, SE of Millaa Millaa, Hale 832386 (US). Stallion Pocket 

logging area, 14 km from Gillies Hwy. and 1 km E from Mulgrave River Forestry rd., S of Gordonvale, Hale 

832388 (US). About 5 km NW of Babinda at the bridge crossing of the Russell River, Hale 831173 (US). 

Francis Range, Woopen Creek Rd, NW of Innisfail, Hale 830623, 830948, 832224 (US). 12.5 km E of 

Cardstone on Tully River Rd. to Kareeya Power Station, W of Tully, Hale 830959, 831726 (US). 11.5 km on the 

Kirrama Forest Rd., W of Kennedy, Hale 831341 (US). About 7.5 km E of Wallaman Falls, W of Ingham, Hale 

832387 (US). Eungella NP., Finch Hatton Gorge, Lumbsch & Mangold 19116 n (F). Sri Lanka, Hale 51157 

(US).


Chapsa halei Mangold spec. nov. ined. 

Type: Australia, Queensland, Mt. Lewis Rd. 12km N from Kennedy Hwy., W of Mossman, Hale 831320 

(US-holotype). 

ETYMOLOGY – This species is named in honor of Mason E. Hale who vastly contributed to 

the knowledge of Thelotremataceae and who left behind a large collection of this family from 

Australia. 


Fig. 10. Chapsa halei: growth habit (A), ascomata (B) and ascospores (C, D). A.-D.: US-holotype. 

Bar= A: 1 mm; B: 0.7 mm; C: 5 µm; D: 3 µm. 

Thallus hypo- to epiphloedal, (moderately) thin, up to c. 150 µm high, pale gray. Surface 

dull, porous to roughened, continuous to rugose, predominantly unfissured. True cortex 

absent, thallus covered by a thin, incontinuous protocortex, up to c. 10 µm thick. Algal layer 

±continuous, moderately well developed, calcium oxalate crystals abundant, usually small 

and scattered to more rarely clustered. Vegetative propagules not seen. Ascomata 

inconspicuous, large, up to c. 800 µm in diam., roundish to more often irregular to somewhat 

elongated, apothecioid when young, becoming chroodiscoid at maturity, erumpent, 

predominantly solitary, immersed. Disc partly to rarely entirely visible from surface, offwhite 

to pale brownish, epruinose to indistinctly pruinose. Proper exciple not visible from 

surface, thalline rim margin irregular, coarsely split, usually with 2-4 large, thick, irregular


lobes, rarely somewhat eroded, concolorous with thallus, becoming erect to recurved. Exciple 

fused, thin, often with substrate layers and crystals incorporated, hyaline internally to pale 

yellowish or yellowish-brown marginally, non-amyloid. Hymenium up to c. 70 µm high, noninspersed, 

weakly conglutinated, paraphyses thick, straight, parallel, unbranched, tips not 

thickened to slightly thickened, lateral paraphyses present, conspicuous, up to c. 20 µm long, 

thick and adspersed with fine granules. Epihymenium thin to indistinct, hyaline, sometimes 

with fine grayish-brown granules. Asci 8-spored, tholus moderately thin, mostly not visible at 

maturity. Ascospores very small, transversely septate, cell walls moderately thick, nonhalonate, 

hyaline, non-amyloid, predominantly fusi- to claviform, with roundish to subacute 

ends, loci roundish, subglobose to oblong to irregular, end cells cone-shaped, divided by 

regular, moderately thin to moderately thick septae, 10-13 x 2-3 µm with 3-6 loci. Pycnidia 

not seen. 


ECOLOGY AND DISTRIBUTION – Chapsa halei 

was collected in Australia on tree bark in a tropical 

rainforest at 800 m. It is only known from the type 

collection in Northern Queensland. 


gray thallus containing numerous crystals, simple, 

coarsely lobed ascoma margins, epruinose to 

weakly pruinose, and bright apothecial discs. 

Anatomical characters of the taxon include thick 

paraphyses and lateral paraphyses, the latter with 

fine granules, and small, narrow, transversely 

septate, hyaline, non-amyloid ascospores with 

moderately thickened parts. The species lacks 

secondary metabolites. A similar Australian species 

is C. lassae, see under this species for differences. 

Another similar Australian species with a thin, 

ecorticate thallus, lacking secondary compounds is 

C. alborosella. This species can be readily 

distinguished by larger ascospores (up to 25 µm 

long, with up to 9 loci). An additional similar taxon is C. dissuta 1 from Panama that differs in 

ascospore morphology. The ascospores in C. dissuta have angular loci that become fused 

centrally but otherwise have thick septae and endospore. This type of ascospores is 

reminiscent to ascospores in the O. clandestina-group (Frisch, 2006: 334). 

SPECIMENS EXAMINED – See type collection. 

Chapsa indica Massal. 


C. halei. 

Atti Reale Ist. Veneto Sci. Lett. Arti, ser. 3, 5: 257 (1860). Type: Sri Lanka, sine loco (VER-holotype). 

Thelotrema pycnophragmium Nyl., Sertum lichenum tropicale Labuan et Singapore: 5 (1891). Ocellularia 

pycnophragmia (Nyl.) Zahlbr., Cat. Lich. Univ. II: 599 (1923). Type: Malaysia, Labuan, 1879, Almquist s.n. (H- 

Nyl.22679!-lectotype, selected by Frisch [2006: 100]). 

1 Chapsa dissuta (Hale) Mangold comb. nov. ined. – Bas.: Ocellularia dissuta Hale, Smithson. Contrib. Bot. 38: 

20 (1978) – Thelotrema dissutum (Hale) Hale, Mycotaxon 11: 131 (1980). Type: Panama, Colon, Santa Rita 

Ridge, Hale 43520 (US!-holotype).


Thelotrema albescens Vain., Boletim Soc. Broteriana, sér. 2, 6: 152 (1929). Ocellularia albescens (Vain.) 

Zahlbr., Cat. Lich. Univ. X: 211 (1939). Type: Mozambique, Palma, 1916, A. Pires de Lima 524, (TUR- 

Vain.34793-lectotype, selected by Salisbury [1971a: 273]). 


Fig. 12. Chapsa indica: ascomata (A), ascus (C) and ascospores (B). A.: Lumbsch & Streimann 

27576; B., C.: H-lectotype. Bar= A: 1 mm; B: 12.5 µm; C: 10 µm. 

Thallus predominantly hypophloedal, very thin, up to c. 50 µm high, pale gray to pale 

grayish-green. Surface dull to slightly shiny, smooth to roughened, often due to protuberant 

substrate, continuous, predominantly unfissured, sometimes appearing fissured due to 

substrate structure. Cortex structures absent to rarely covered by a thin, incontinuous 

protocortex up to c. 10 µm thick. Algal layer poorly to moderately well developed, continuous 

to incontinuous, calcium oxalate usually sparse, small to moderately large, scattered to more 

rarely clustered. Vegetative propagules not seen. Ascomata variable, conspicuous to 

inconspicuous, (moderately) large, up to c. 1.3 mm in diam., roundish to irregular, fused 

ascomata sometimes appear slightly branched, chroodiscoid at mature stages, erumpent, 

solitary to fused, immersed. Disc partly to entirely visible from surface, grayish, usually 

distinctly pruinose, sometimes glittering. Proper exciple not visible from surface to partly 

visible when becoming free, thalline rim split, distinctly lobed to more rarely somewhat 

eroded, lobes large and thick, internally ±pruinose, whitish to somewhat brownish from 

protuberant substrate, concolorous with thallus outside, erect to recurved. Exciple fused to 

partly free, thin to evanescent, colorless internally to yellowish or pale orange and often with 

substrate layers incorporated marginally, apically often covered with grayish granules, nonamyloid. 

Hymenium up to c. 150 µm high, non-inspersed, weakly conglutinated, paraphyses 

±straight, parallel to slightly interwoven, unbranched, tips moderately to distinctly thickened, 

lateral paraphyses present, inconspicuous, up to c. 20 µm long. Epihymenium (moderately)


thick, with grayish granules, sometimes with small crystals. Asci 6 to 8-spored, tholus thick, 

remaining thickened at maturity. Ascospores (moderately) large, transversely septate, cell 

walls (moderately) thick, endospore moderately thin to moderately thick, non-halonate, 

hyaline, non-amyloid, oblong-fusiform, with narrowed-roundish to subacute ends, loci 

roundish to slightly angular, rarely subglobose to lentiform towards the ends to predominantly 

oblong, end cells hemispherical to conical, septae (moderately) thin, regular, 50-110 x 6-12 

µm with 20-35 loci. Pycnidia not seen. 


ECOLOGY AND DISTRIBUTION – Chapsa indica 

was collected in Australia on the bark of an 

undetermined tree in a tropical rainforest in an 

altitude of 60 m. It is an extremely rare species in 

Australia, only known from north-western Northern 

Territory. This is the first report for Australia and 

the Philippines. It has been recorded from Africa, 

India, Sri Lanka, Andaman Islands and Malaysia 

(for references see Frisch, 2006) indicating a 

paleotropical distribution. 

N OTES – This species is characterized by the 

very thin, ecorticate thallus, the whitish-pruinose 

ascomata, the large, transversely septate, hyaline, 

non-amyloid, oblong-fusiform ascospores with 

thickened cell-walls and the lack of lichen 

substances. The most similar species in Australia is 

C. pulchra, which differs by its ascospore 

morphology. It has smaller (up to 70 µm long, with 

up to 24 loci), thin-walled, predominantly 

cylindrical ascospores that are fragile and easily break apart. 

SPECIMENS EXAMINED – Australia, Northern Territory: Curtain Falls, Litchfield Park, 38km WSW of 

Batchelor, Elix, Lumbsch & Streimann 27576 (CANB). Philippines, Mindanao, Agusan, Nasipit Lumber Co., 

Florida logging area, c. 30 km SE of Butuan, Hale & Banaag 25426 (US). 

Chapsa lamellifera (Kantvilas & Vezda) Mangold comb. nov. ined. 

Bas.: Chroodiscus lamelliferus Kantv. & Vezda, Lichenologist 32: 336 (2000). Type: Australia, Tasmania, 

Ben Ridge, Kantvilas 105/81 (HO!-holotype; hb.Vezda-isotype). 



C. indica. 

Thallus epi- to hypophloedal, thin to very thin, up to c. 100 µm high, grayish-green to pale 

gray. Surface dull to slightly shiny, smooth, continuous, unfissured. Thallus covered by an 

incontinuous protocortex up to c. 20 µm thick. Algal layer ±continuous, moderately well 

developed, calcium oxalate crystals sparse, small, scattered. Vegetative propagules not seen. 

Ascomata variable throughout development, conspicuous, (very) large, up to c. 3 mm in 

diam., predominantly roundish, chroodiscoid at maturity, first erumpent to sessile in older 

stages, solitary to fused, becoming ±distinctly emergent, predominantly depressed-urceolate. 

Disc partly to more rarely entirely visible from surface, grayish-brown to flesh colored,


indistinctly pruinose. Proper exciple not visible from surface, thalline rim characteristic, 

±regularly radially split, lobed, becoming strongly layered, off-white to pale reddish-brown, 

inner layer(s) predominantly incurved, often gradually exfoliating, becoming erect to 

recurved towards the outer layers. Proper exciple fused, moderately thin to somewhat 

evanescent, hyaline internally to pale brownish marginally, non-amyloid. Hymenium up to c. 

180 µm high, non-inspersed, strongly conglutinated, paraphyses straight to slightly bent, 

parallel, unbranched, tips unthickened to slightly thickened, lateral paraphyses present, 

predominantly inconspicuous, up to c. 25 µm long. Epihymenium thin, hyaline with grayishbrown 

granules. Asci 8-spored, tholus absent. Ascospores moderately small to moderately 

large, eumuriform, cell walls thin, with thin halo, hyaline to slightly yellowish in older 

ascospores, non-amyloid, oblong to (irregularly-)ellipsoid to reniform, with rounded to 

narrowed-rounded ends, loci small, roundish to angular, irregular, transverse septae only 

distinct in younger ascospores, irregular, vanishing with age, 30-60 x 10-20 µm with 12-18 x 

2-5 loci. Pycnidia not seen. 

Fig. 14. Chapsa lamellifera: ascomata (A, B) and ascospores (C, D). A.-D.: HO-holotype. Bar= A: 

1.5 mm; B: 3 mm; C: 15 µm; D: 10 µm. 

CHEMISTRY – Thallus K+ yellowish, C-, PD+ reddish; containing protocetraric, 

fumarprotocetraric (majors to minors) and succinprotocetraric (minor) acids. 

ECOLOGY AND DISTRIBUTION – Chapsa lamellifera was collected on the bark of trees, dead 

wood and more rarely on soil, debris or moss in moist, cool temperate rainforests in altitudes 

ranging from sea level to 900 m. It is common in Tasmania and currently only known from 

there and from New Zealand (Kantvilas & Vezda, 2000).


NOTES – This species can be readily identified 

by the large, distinctly layered ascomata, the 

moderately large, muriform, hyaline, thin walled, 

non-amyloid ascospores and the presence of the 

protocetraric acid chemosyndrome. A very close 

species is C. asteliae, which is also known from 

Tasmania. It can be distinguished by the different 

ecology (on Astelia in [sub-]alpine heathlands), 

smaller ascomata with grayish-pruinose discs and 

narrower ascospores with fewer longitudinal septae 

that show a faint amyloid reaction. Also similar, 

but readily distinguished by the transversely septate 

ascospores is C. minor, see also under this species. 

Topeliopsis macrocarpa and T. tasmanica could be 

also confused with C. lamellifera, but can be easily 

differentiated by the larger ascospores and the 

presence of the stictic acid chemosydrome. Vezda 

erroneously distributed C. lamellifera as an 

exsiccate of C. megalophthalma (as Chroodiscus 

megalophthalmus / Lich. Rar. Exsicc. n. 25). The 

latter taxon is another similar species, for differences see under this species. Many of the 

examined collections were sterile, however, they could be identified by their characteristic 

habitus and chemistry. 

SPECIMENS EXAMINED – Australia, Tasmania: King William Saddle, 14 km SW of Derwent Bridge, Elix 

26939 (CANB, B). Elliot Range, (Vezda: Lich. Rar. Exsicc. n. 25, C. megalophthalmus), Kantvilas & Jarman 

s.n., (BM, F, GZU, H, UPS). Southwest Conservation Area, along Scotts Peak Rd., 2.5 km from Gordon River 

Rd. turnoff, at Rainforest Nature Walk, Wedin 3048 (UPS). Lake Pedder, SW Tasmania, Bratt 2777 (BM). 

Uncertain locality: "On a stunted (Notho-)Fagus in an exposed situation alt. 2000 ft.", Gunn 1763 (NSW). 

Chapsa lassae Mangold spec. nov. ined. 

Type: Australia, Queensland, Daintree NP., Mossman Gorge Section, near Rex Creek Swing Bridge, 

Mangold 35 zq (CANB-holotype, BRI-, F-isotypes). 

ETYMOLOGY – This species is named after Anne Lass who was not only of great help on 

the second field trip to Australia, but also supported me throughout the whole project. 



C. lamellifera. 

Thallus hypo- to mainly epiphloedal, in parts bulging and flaking away from the substrate, 

(moderately) thin, up to c. 100 µm high, pale olive. Surface shiny, smooth, continuous, 

unfissured. True cortex present, predominantly thin, ±continuous, consisting of irregular to 

more rarely periclinal hyphae, up to c. 20 µm thick. Algal layer well developed, continuous, 

calcium oxalate crystals lacking. Vegetative propagules not seen. Ascomata conspicuous, 

moderately large, up to c. 600 µm in diam., roundish to more often irregular, apothecioid to 

chroodiscoid in mature ascomata, erumpent, solitary to fused, often somewhat clustered, 

regenerating, immersed. Disc partly to rarely entirely visible from surface, grayish-brown, 

indistinctly pruinose. Proper exciple not visible from surface, thalline rim margin irregular, 

coarsely split, ±irregularly lobed, becoming distinctly layered in subsequent ascomata 

generations, off-white to pale reddish-brown, incurved to erect, outer layers becoming 

recurved. Exciple fused, (moderately) thin, with substrate layers incorporated, hyaline


Fig. 16. Chapsa lassae: growth habit (A), ascomata (B), ascoma section (C) and ascospores (D, E). 

A.-E.: CANB-holotype. Bar= A: 1 mm; B: 0.5 mm; C: 100 µm; D, E: 3 µm. 

internally to pale yellowish or yellowish-brown marginally, non-amyloid. Hymenium up to c. 

80 µm high, non-inspersed, weakly conglutinated, paraphyses ±straight, moderately 

interwoven, unbranched, tips slightly to distinctly thickened, irregular, lateral paraphyses 

present, inconspicuous, up to c. 25 µm long. 

Epihymenium thin, hyaline, with fine grayishbrown 

granules. Asci 8-spored, tholus moderately 

thick, becoming (moderately) thin at maturity. 

Ascospores typical, very small, transversely 

septate, cell walls and endospore thin, with distinct, 

thick halo, hyaline, non-amyloid, predominantly 

fusi- to claviform, with roundish to subacute ends, 

loci large, angular, end cells hemispherical to 

conical, divided by ±regular, thin septae, 10-15 x 4- 

6 µm with 3-5 loci. Pycnidia not seen. 

CHEMISTRY – Thallus K-, C-, PD-; no 

compounds detectable by TLC. 

ECOLOGY AND DISTRIBUTION – This species was 

collected on tree bark in a tropical rainforest at 

100 m altitude. It is only known from the type 

collection in northern Queensland. 


C. lassae.


NOTES – Chapsa lassae is a characteristic species with a shiny, thin, but mainly 

epiphloedal thallus, regenerating ascomata with reddish-brown, layered margins, slightly 

pruinose, grayish-brown discs, and small, transversely septate, hyaline, non-amyloid 

ascospores with thin call walls and septae and a distinct, thick halo. Similar species in 

Australia include C. alborosella, C. astroidea and C. halei. The latter species, as well as the 

also similar C. dissuta from Panama can be distinguished be the absence of layered ascomata 

margins and the non-halonate ascospores with distinctly thickened parts. For differences to 

C. alborosella and C. astroidea see under this species. 


Chapsa leprieurii (Mont.) A. Frisch 

Bibl. Lich. 92: 105 (2006). Bas.: Stictis leprieurii Mont., Ann. Sci. Nat., Bot., sér. 4, 3: 97 (1855). 

Phaeotrema leprieurii (Mont.) Sherwood, Mycotaxon 5: 203 (1977). Thelotrema leprieurii (Mont.) Hale, 

Mycotaxon 11: 131 (1980). Type: French Guiana, Cayenne, Leprieur 804 (PC-lectotype, selected by Hale [1981: 

258]; G-isolectotype) 

Thelotrema leucastrum Tuck., Proc. Amer. Acad. Arts Sci. 6: 269 (1864). Phaeotrema leucastrum (Tuck.) 

Zahlbr., Cat. Lich. Univ. II: 608 (1923). Type: Cuba, Wright, Lichenes Cubae 158 (FH-Tuck.-lectotype, selected 

by Frisch [2006: 105]; H-Nyl. 22662!-, L-, M!-isolectotypes). 

Thelotrema leucastrum var. difforme Tuck., Proc. Amer. Acad. Arts Sci. . 6: 269 (1864). Thelotrema 

difforme (Tuck.) Vain., Ann. Acad. Sci. Fenn. A 15(6): 194 (1921a). Phaeotrema leucastrum var. difforme 

(Tuck.) Zahlbr., Cat. Lich. Univ. II: 608 (1923).Type: Cuba, Wright, Lichenes Cubae 159 (FH-Tuck.-holotype; 

H-Nyl. 22664-, L-, M!- isotypes). 

Graphis subnivescens Nyl., Boletim. Soc. Broteriana 4: 211 (1886a); Flora 69: 174 (1886b). Phaeotrema 

subnivescens (Nyl.) Zahlbr., Cat. Lich. Univ. II: 387 (1923). Type: São Thomé, 1885, Moller s.n. (H-Nyl. 7507holotype). 

Graphis phlyctidea Vain., Ann. Acad. Sci. Fenn. A 15(6): 137 (1921a). Phaeographis phlyctidea (Vain.) 

Zahlbr., Cat. Lich. Univ. II: 383 (1923). Type: Philippines, Luzon, Irosin, Elmer 14646 (TUR-Vain. 27523holotype; 

FH-isotype). 

Thelotrema confluens Vain., Ann. Acad. Sci. Fenn. A 15(6): 193 (1921a) nom. illegit. [non Thelotrema 

confluens Kremp.]. Ocellularia confluentula Zahlbr., Cat. Lich. Univ. II: 587 (1923) nom. nov. pro Thelotrema 

confluens Vain. Type: Philippines, Luzon, Irosin, Elmer 14641 (pr. p.)/14623 (TUR-Vain.-26908!-holotype). 


Thallus hypo- to epiphloedal, (moderately) thin, up to c. 300 µm high, dark olive to olivebrown 

or pale yellowish-brown. Surface dull to wax-like, smooth, continuous to slightly 

verrucose, unfissured. True cortex present, ±continuous, up to c. 30 µm thick, pale yellowish, 

consisting of periclinal to irregular hyphae. Algal layer continuous to incontinuous and 

moderately well developed, calcium oxalate crystals usually abundant, mostly small and 

scattered, sometimes clustered. Vegetative propagules not seen. Ascomata very conspicuous, 

(moderately) large to rarely very large, up to c. 1.7(2) mm in diam., roundish to irregular, 

particularly in fused ascomata, apothecioid to chroodiscoid in older ascomata, erumpent, 

solitary to fused, often clustered in groups of two to several ascomata, immersed. Disc 

sometimes not visible to more often partly or rarely entirely visible from surface, grayish, 

distinctly pruinose, sometimes covered with same whitish-cottony substance as thalline rim. 

Proper exciple not visible from surface to more rarely becoming visible when partly detached, 

whitish, thalline rim (moderately) thick, indistinctly split to entire, often appearing somewhat 

eroded, with a ±thick whitish, cottony surface with sometimes visible tiny crystal needles, 

predominantly erect to recurved. Exciple fused to apically partly free, thin, colorless to pale 

yellowish internally to (pale) orange or brownish marginally, usually apically densely covered 

by grayish granules. Hymenium up to 110 µm high, non-inspersed, moderately conglutinated,


paraphyses straight to slightly bent, parallel to slightly interwoven, unbranched, tips 

±distinctly thickened, slightly irregular, lateral paraphyses present, conspicuous, up to 35 µm 

long. Epihymenium (moderately) thick, hyaline, with grayish granules. Asci 8-spored, tholus 

moderately thick, becoming thin or not visible at maturity. Ascospores (very) small, 

transversely septate, rarely with a singular longitudinal septum, cell walls and endospore 

(moderately) thick, non-halonate, brown, distinctly amyloid, oblong to ellipsoid to somewhat 

clavate with rounded to narrowed-rounded to rarely subacute ends, loci roundish, subglobular 

to more often lentiform or oblong, end cells hemispherical to more rarely conical, septae 

moderately thin to moderately thick, regular at maturity, 9-25(28) x 6-8(11) µm with 4- 

7(8)x(2) loci. Pycnidia not seen. 


Fig. 18. Chapsa leprieurii: growth habit (A), ascomata (B, C) and ascospores (D-F). A., D.-F.: Hale 

830708; B.: Lumbsch & Mangold 19127 h; C.: H-isolectotype of T. leucastrum. Bar= A: 1.2 mm; 

B: 0.5 mm; C: 0.8 mm; D: 6 µm; E: 5 µm; F: 4 µm.



leprieurii was collected in Australia on tree bark in 


1000 m. It is regionally common in northern 

Queensland. This is the first report for Australia, 

outside of Australia it is known from the 

Neotropics (Hale, 1981; Frisch, 2006), Africa 

(Hale, 1981; Frisch, 2006), India (Awasthi, 1991), 

Sri Lanka Sumatra and Philippines (Hale, 1981), 

indicating a pantropical distribution. 

NOTES – This species is characterized by a thin, 

corticate, dark thallus that gives a conspicuous 

contrast to the bright ascomata that have a typical 

cottony surface structure, at least in freshly 

collected material. Further it has small, transversely 

septate (a single longitudinal septum may rarely 

occur), brown, amyloid ascospores with distinctly 

thickened parts, and it lacks secondary compounds. 

In the examined specimens the size of the ascomata 

varied remarkably. Whereas most of the apothecia are indistinctly chroodiscoid and do not 

exceed 1 to 1.5 mm in diameter, a few ascomata in the type material of T. leucastrum show a 

typical chroodiscoid habitus and are up to 2 mm in diameter (see also Fig. 18., B and C). In 

Australia, a similar species is the stictic acid containing C. platycarpa, which further differs in 

slightly smaller (up to 20 µm long, with up to 6 loci), never distinctly amyloid ascospores and 

a distinctly free proper exciple. Another similar species that lacks lichen substances is C. 

aggregatum, known from Dominica and Sri Lanka. It differs by the distinctly free proper 

exciple, the usually apothecioid ascomata and smaller ascospores (up to 18 µm long, with up 

to 6 loci). 

SPECIMENS EXAMINED – Australia, Queensland: Iron Range NP., 29 km from western boundary on track to 

Portland Rds., Hale 830058 (US). Cape Tribulation, 2 km W of main rd. between Oil Palms and Coopers Creek, 

N of Daintree, Hale 832163 (US). Thornton Range, CREB rd. (to Cooktown), about 5 km in from Daintree 

River crossing, NW of Mossman, Hale 831398 (US). 15 km along Mulgrave River Rd. from intersection with 

the Gordonvale Rd., SW of Gordonvale, Hale 830725, 830683, 830686 (US). Stallion Pocket logging area, 14 

km from Gillies Hwy and 1 km E from Mulgrave River Forestry rd., S of Gordonvale, Hale 832752, 832706 

(US). Atherton Tablelands: Crystal Cascades, Lumbsch & Mangold 19117p (F); Lake Tinaroo, Lumbsch & 

Mangold 19127h, Mangold 38i (F). The Boulders, NW of Babinda, S of Cairns, Hale 831350 (US). Bellenden 

Ker NP., Boulders area, 6 km W of Babinda, 50 km S of Cairns, A. & M. Aptroot 22412 (ABL). Francis Range, 

Woopen Creek Rd., 25 km in from Bruce Hwy., NW of Innisfail, Hale 832156, 832384, 832793 (US). Mt. 

Chalmynia logging area, 15 km from Bruce Hwy., W of Innisfail, Hale 832071, 832367, 832385 (US). 

Josephine Falls, W of Bartle Frere and Bruce Hwy., S of Innisfail, Hale 830706 (US). Mt. Spec NP., Ridge on 

the Loop, on the Paluma Rd., WNW of Townsville, Hale 830708 (US). 

Chapsa leprocarpa (Nyl.) A. Frisch 

Bibl. Lich. 92: 108 (2006). Bas.: Graphis leprocarpa Nyl., Acta Soc. Sci. Fenn. 7: 472 (1863a). Thelotrema 

leprocarpum (Nyl.) Tuck., Genera lichenum: 139 (1872). Graphina leprocarpa (Nyl.) Zahlbr., Cat. Lich. Univ. 

II: 412 (1923). Type: U.S.A., Louisiana, 1853, Hale 111 (FH-Tuck.-holotype; H-Nyl. 6839!-isotype). 

Thelotrema colobicum Nyl., Bull. Soc. Linn. Normandie, sér. 2, 7: 169 (1874 ['1873']). Type: Andaman 

Islands, 1867, Kurz 43 (M-holotype; H-Nyl.22493!-isotype). 



C. leprieurii.


Fig. 20. Chapsa leprocarpa: growth habit (A) and ascospore (B). A., B.: Mangold 37 h. Bar= A: 0.75 

mm; B: 17 µm. 

Thallus predominantly hypophloedal, very thin to thin, up to c. 70 µm high, pale gray to 

pale grayish-green, on dark substrate accordingly darker. Surface dull, rough to somewhat 

pruinose, often with protuberant substrate structure, continuous, unfissured. Cortex structures 

absent. Algal layer poorly developed, predominantly incontinuous, calcium oxalate crystals 

lacking to moderately abundant, usually small, clustered or scattered. Vegetative propagules 

not seen. Ascomata conspicuous, (moderately) large, up to c. 1.7 mm in diam., but more often 

not larger than c. 700 µm in diam., roundish to slightly irregular, particularly in fused 

ascomata, apothecioid to chroodiscoid in older ascomata, erumpent, solitary to fused, 

sometimes regenerating, immersed. Disc partly to entirely visible from surface, grayish, 

usually distinctly pruinose. Proper exciple not visible from surface to visible when becoming 

partly detached, whitish, thalline rim margin irregular, usually coarsely split with large lobes, 

sometimes eroded, slightly to more rarely distinctly layered, whitish-pruinose internally, 

concolorous with thallus marginally, becoming erect to ±recurved. Proper exciple thin to 

somewhat evanescent, hyaline internally to pale yellowish to pale orange marginally, usually 

with substrate layers incorporated, apical parts covered by grayish granules, slightly to 

distinctly amyloid at the base. Hymenium up to c. 140 µm high, non-inspersed, weakly 

conglutinated, paraphyses straight to slightly bent, parallel to slightly interwoven, 

unbranched, tips distinctly thickened, lateral paraphyses present, conspicuous, up to c. 30 µm 

long, adspersed with fine granules. Epihymenium predominantly thick, hyaline, with coarse 

grayish-brown granules. Asci 1- to more rarely 2-spored, tholus thin, not visible at maturity. 

Ascospores (moderately) large, densely eumuriform, cell walls and endospore thin, with a 

thin and indistinct to a thick, distinct halo, then strongly thickened at the ascospore ends, 

hyaline, non-amyloid, oblong to slightly fusiform, with roundish to narrowed-roundish ends, 

loci small, angular to slightly roundish, transverse septae distinct, thin, regular, 60-130 x 20- 

40 µm µm with multiple loci. Pycnidia not seen. 

CHEMISTRY – Thallus K-, C-, PD-; no compounds detectable by TLC.



leprocarpa was collected in Australia from tree 

bark in (sub)tropical rainforests in altitudes ranging 

from sea level to 1100 m. It is a common species, 

occurring in northern Queensland and in the wider 

Queensland/ New South Wales border region. This 

is the first report for Australia, otherwise it is 

known from the Neotropics (Frisch, 2006; 

Tuckerman, 1872), Africa (Frisch, 2006), India 

(Patwardhan & Nagarkar, 1980), Sri Lanka (Hale, 

1981) and Borneo (Sipman, 1993) indicating a 

pan(sub)tropical distribution. 


ecorticate thallus, often indistinctly chroodiscoid 

ascomata with a whitish-pruinose surface, conspicuous 

lateral paraphyses with fine granules, 

large, densely eumuriform, hyaline, non-amyloid 

ascospores with thin cell walls that occur singular 

ore rarely by two per ascus and the lack of lichen 


C. leprocarpa. 

substances. In Australia, C. niveocarpa is a similar species see under this species for 

differences. On the world level, there are two species that can be confused with C. 

leprocarpa, the very similar C. patens that can be distinguished by larger ascospores (up to 

160 µm long), and C. grossomarginata 2 that has smaller ascospores (up to 80 µm long) with 

less numerous, larger loci, and lateral paraphyses without granular structures. 

SPECIMENS EXAMINED – Australia, Queensland: Big Tableland, 26 km S of Cooktown, Streimann 30901 

(CBG). 14 km SW of Mossman, Mt. Lewis, Tibell 14598 (UPS). Near end of Black Mountain Rd., 33 km WNW 

of Kuranda, Hale 832079 (US). Fresh Water Gorge, outside of Cairns, Hale 832487 (US). Rd. from Gordonvale 

to Yarrabah, c. 10 km E of Cairns, Lumbsch & Guderley 11158 u (F). Atherton Tablelands: Danbulla Forest 

Drive, 4 km E of Tinaroo, Hale 831212 (US); Lake Tinaroo, Downfall Creek Camping Area, Lumbsch & 

Mangold 19125 k (F); Bunbulla Forest Drive, near Lake Euramoo, Mangold 37 h (F); Lake Eacham NP., 

Mangold 29 c, be (F); 20 km SE of Yungburra, 1 km NW of Bonjes, Tibell 15343 (UPS); Area below crater, Mt. 

Hypipamee NP., S of Atherton, Hale 831924 (US); Malanda Falls, just N of Malanda, Lumbsch & Mangold 

19132 m (F); SW of K-1 tree rd. off Palmerston Hwy., 11 km from main hwy. and 2 km N of S. Johnstone 

Forestry Camp, SE of Millaa Millaa, Hale 832182 (US); Tumoulin Rd., 5 km from turnoff to Ravenshoe, 

Lumbsch & Mangold 19133 u, 19136 g, 19151 q, Mangold 30 zi, zm (F); Just S of hwy., 13 km E of jct. 

Kennedy Hwy. and Palmerston Hwy., E of Ravenshoe, Hale 831420, 832131 (US). State Forest area on Tully 

Rd., 1 km from jct. with S. Mission Beach Rd., S of Mission Beach, Hale 831540 (US). Mt. Spec NP., Ridge on 

the Loop, on the Paluma Rd., WNW of Townsville, Hale 831856, 832322, 832390 (US). Eungella NP.: NP. side 

rd. near Pease's Lookout, off Darymple Rd., Hale 831413 (US); Finch Hatton Gorge, Lumbsch & Mangold 

19113 u (F); Trail at Broken River Picnic Area, Hale 831679 (US). Noosa NP.: Palm Grove Track, Hale 831452 

(US); c.1 km from the sea near rd. to the Lookout, Thor 4902/II (S). Noosa Heads, K. & A. Kalb 34277 (hb. 

Kalb). Wooroi State Forest Park, W of Teewantin, Hale 832756 (US). Sunshine Coast Hinterland, Kenilworth 

SF., Lumbsch & Mangold 19085 e (F). New South Wales: Nightcap Forest Drive: Gibbergunyah Roadside 

Reserve, Whian Whian SF., W of Mullumbimby, Hale 831505 (US); 1 km W of Minyon Falls, N of Lismore, 

Hale 832215 (US). Tweed Range, Mebbin NP., Mangold 21 f, i (F). Dorrigo NP., Sassafras Creek Track, 

Mangold 25 d, p (F). 

2 Chapsa grossomarginata (Matsum.) Mangold comb. nov. ined. Bas.: Thelotrema grossomarginatum Matsum., 

J. Hattori Bot. Lab. 88: 20 (2000). Type: Japan, Honshu, Matsumoto 2492 (HIRO!-holotype).


Chapsa lordhowensis Mangold spec. nov. ined. 

Type: Australia, New South Wales, Lord Howe Island, Goat House Cave, Elix 42259 (CANB-holotype). 

ETYMOLOGY – The epithet refers to the type location Lord Howe Island. 


Fig. 22. Chapsa lordhowensis: growth habit (A), ascomata (B, C), ascospore (D), ascospore with 

amyloid reaction (E), pycnidia (F) and conidia (G). A., D., E.: Elix 42267; B., F., G.: CANBholotype; 

C.: Elix 42259. Bar= A: 2.5 mm; B: 2 mm; C: 1 mm; D: 12.5 µm; E: 14 µm; F: 300 µm; 

G: 5 µm.


Thallus predominantly epiphloedal, moderately thick, up to c. 300 µm high, rarely pale 

yellowish-brown to usually pale olive. Surface ±shiny, smooth, continuous, unfissured. True 

cortex present, thick, continuous, up to c. 50 µm thick, hyaline to slightly yellowish, 

consisting of periclinal to irregular hyphae. Algal layer continuous and well developed, 

calcium oxalate crystals sparse, sometimes clusters of small crystals are found in lower 

thallus. Vegetative propagules not seen. Ascomata variable, conspicuous, moderately large to 

very large, up to c. 3 mm in diam., roundish to more rarely somewhat irregular, sometimes 

appearing slightly branched in fused ascomata, apo- to chroodiscoid, erumpent, solitary to 

rarely fused, typically regenerating, ±emergent, hemispherical to urceolate in younger 

ascomata, depressed-urceolate to cupular in older stages. Disc variable, partly to entirely 

visible from surface, pale to distinctly flesh-colored, epruinose to slightly pruinose, in 

deceasing ascomata disc becoming overgrown by a off-white to pale yellowish-brown layer. 

True exciple not visible from surface, thalline rim margin variable throughout ontogeny, in 

newly developing ascomata split to rugged or lobed, ±concolorous with thallus or brighter, 

becoming erect to more rarely recurved, in subsequent ascomata generations, margin 

developing out of layer covering the disc, splitting and opening irregularly, breaking away in 

large parts, marginally parts finally forming the new thalline rim, becoming erect to rarely 

recurved, layered with age, off-white to pale yellowish-brown. Exciple fused, moderately 

thick, hyaline internally to pale yellowish or yellowish-orange marginally, usually exciple 

extending into a distinct hyaline internal exciple moderately to strongly amyloid. Hymenium 

up to c. 180 µm high, non-inspersed, moderately conglutinated, paraphyses slightly bent, 

parallel, unbranched, tips predominantly unthickened, lateral paraphyses present, 

inconspicuous, up to c. 25 µm long. Epihymenium thick, hyaline to pale yellowish, with 

grayish- to yellowish-brown granules. Asci 1- to rarely 2-spored, tholus moderately thick, not 

visible at maturity. Ascospores (moderately) large, densely eumuriform, cell walls 

(moderately) thin, endospore thin, non-halonate, hyaline, distinctly amyloid, oblong to 

ellipsoid with rounded to narrowed-rounded ends, loci roundish to angular, predominantly 

irregular, distinctly transversely divided only in younger stages, in older stages septae 

becoming irregular and finally vanish, 65-120 x 22-35 µm with multiple loci. Pycnidia 

present, conspicuous, variable, predominantly in strongly raised verrucae with unconstricted 

to strongly constricted to somewhat stalked base and usually with bright tip becoming 

blackish when mature, Conidia small, irregular-roundish to irregular-oblong, up to c. 2 x 1 

µm in size. 

CHEMISTRY – Thallus K+ yellowish to brown, 

C-, PD+ orange; containing stictic, constictic 

(majors), α-acetyloconstictic, cryptostictic, 

α-acetylhypoconstictic (minors), hypoconstictic 

and hypostictic (traces) acids. 

ECOLOGY AND DISTRIBUTION – Chapsa lordhowensis 

was collected in Australia on tree bark in 

subtropical rainforests in mid-range altitudes of 

380-420 m. It is only known from the typecollection 

on Lord Howe Island and from north of 

Sydney in continental New South Wales. 

NOTES – The taxon is characterized by a shiny, 

distinctly corticate thallus, regenerating ascomata 

with rugged, layered margins, flesh-colored discs 

that become covered by a bright layer, large, 


C. lordhowensis.


hyaline, densely eumuriform, thin-walled, amyloid ascospores that occur mostly singular per 

ascus and the stictic acid chemosydrome. Chapsa megalophthalma is a similar, stictic acid 

containing species which can be readily distinguished by 8-spored asci, smaller ascospores 

(up to 45 µm long), and more distinctly Geaster-like ascomata with a less distinctly layered 

margin. Chapsa lamellifera is another similar species that also differs in 8-spored asci with 

smaller, non-amyloid ascospores (up to 55 µm long) and a different chemistry (protocetraric 

acid). This is the first report of pycnidia in Chapsa, the conidia resemble the type found in 

Thelotrema, Acanthotrema and Leucodecton. 

SPECIMENS EXAMINED – Australia, New South Wales: Benowie Walking Track (NNW of Sydney), Archer 

418415 (NSW). Lord Howe Island, Track to Goat House Cave, Elix 42267 (B), 42144, 42146, 42259, 42267 

pr.p. (CANB). 

Chapsa megalophthalma (Müll. Arg.) Mangold comb. nov. ined. 

Bas.: Thelotrema megalophthalmum Müll. Arg., Flora 65: 500 (1882). Chroodiscus megalophthalmus (Müll. 

Arg.) Vezda & Kantv. in Vezda, Lich. Rar. Crit. Exsicc. 3: 2 (1992). Type: Australia, Queensland, Toowoomba, 

Hartmann s.n. (BM!-lectotype, selected by Hale [1972, in herb.]). 

Thelotrema leucophthalmum Nyl. var. lacerata Räs., Suom. Elain-ja Kasvit. Seuran Van. Tiedon. Pöytäkirjat 

3: 184 (1949). Type: Australia, New South Wales, Katoomba, 1889, Wilson s.n. (H!-holotype). 


Fig. 24. Chapsa megalophthalma: growth habit (A), ascomata (B), hymenium and asci (C) and 

ascospore (D). A.-D.: Mangold 19 zb. Bar= A: 2.5 mm; B: 1.5 mm; C: 50 µm; D: 6 µm.


Thallus corticolous to rarely saxicolous, predominantly episubstratic, moderately thick, up 

to c. 200 µm high, rarely pale gray to usually pale grayish-green or (pale) olive. Surface rarely 

dull to ±shiny, smooth, continuous to moderately verrucose or verruculose, unfissured. 

Thallus covered by an incontinuous to continuous protocortex, sometimes becoming 

somewhat conglutinated forming a true cortex of irregular to more rarely periclinal hyphae, 

up to c. 30 µm thick. Algal layer continuous and well developed, calcium oxalate crystals 

lacking. Vegetative propagules not seen. Ascomata conspicuous, (very) large, up to c. 3 mm 

in diam., round to slightly irregular, sometimes appearing slightly branched in fused 

ascomata, chroodiscoid, erumpent, solitary to fused, regenerating, immersed. Disc partly to 

more often entirely visible from surface, variable, dark grayish-brown to brownish to pale or 

distinctly flesh-colored, epruinose to slightly pruinose, in deceasing ascomata disc becoming 

overgrown by a off-white to pale yellowish-brown covering tissue. True exciple superficially 

not distinguishable, thalline rim margin structure variable throughout ontogeny, in newly 

developing ascomata split, rugged to lobed, ±concolorous with thallus or brighter, becoming 

erect to recurved, in subsequent ascomata generations, margin developing out of a tissue 

covering the disc, the tissue splitting and opening irregularly, breaking away in large parts, 

marginal parts finally forming the new thalline rim, off-white to pale yellowish-brown, 

becoming erect to recurved, layered with age (non-regenerating, deceased ascomata 

sometimes also becoming entirely covered by thallus, with remnants of the margin often 

remaining visible). Exciple fused, thin, hyaline internally to pale yellowish or yellowishorange 

marginally, usually lower exciple extending into a distinct hyaline, moderately to 

strongly amyloid lateral hypothecium. Hymenium up to c. 180 µm high, non-inspersed, 

moderately conglutinated, paraphyses ±straight, ±parallel, unbranched, tips unthickened to 

slightly thickened, lateral paraphyses present, inconspicuous, up to c. 20 µm long. 

Epihymenium moderately thin, (pale) brownish, with yellowish-brown to brownish granules. 

Asci 8-spored, tholus thin, not visible at maturity. Ascospores moderately small, eumuriform, 

cell walls (moderately) thin, endospore very thin, non-halonate, hyaline, non-amyloid, oblong 

to ellipsoid to reniform with rounded to more rarely narrowed-rounded ends, loci large, 

roundish to angular, ±rectangular to irregular, transverse septae thin, regular, 25-45(50) x 8- 

12 µm with 8-13(15) x 1-6 loci. Pycnidia not seen. 


C-, PD+ orange; containing stictic (major), 

constictic (major to minor), α-acetylhypoconstictic, 

cryptostictic, α-acetylconstictic, hyposalazinic and 

hypostictic (traces) acids. 


megalophthalma was collected in Australia on tree 

bark and on rocks in shaded habitats in temperate 

rainforests and more rarely wet sclerophyll forests 

in predominantly higher altitudes ranging from 400 

to 1250 m. It is a moderately common species 

occurring throughout New South Wales (except the 

type collection from Queensland near the New 

South Wales border). Besides Australia it was so 

far only reported from New Zealand (Kantvilas & 

Vezda, 2000). Fig. 25. Australian distribution of 

C. megalophthalma.


NOTES – The species is quite unique by having a smooth, often distinctly green thallus, 

large, regenerating ascomata with bright, off-white rimmed, flesh-colored discs. Two similar, 

Australian species are C. lamellifera and C. lordhowensis. Both species can be distinguished 

by a stronger layered ascoma margin and larger ascospores (up to 120 µm long in C. 

lordhowensis / up to 60 µm long in C. lamellifera). Chapsa lamellifera also differs in 

containing the protocetraric acid chemosyndrome. One collection from Cairns (Trinity Bay) 

made by Sayer in 1886 (MEL-26236), determined as T. magalophthalmum is sterile and could 

not be verified. It contains the stictic acid chemosyndrome. However, its identification is 

doubtful since it has large calcium oxalate crystals and strongly thickened paraphyse tips. 

Chapsa recurva from Philippines and India is a similar stictic acid containing species, which 

differs by a distinctly pruinose disc, larger ascospores (up to 68 µm long) and 2-4-spored asci. 

SPECIMENS EXAMINED – Australia, New South Wales: Mt. Warning NP., track from summit to parking lot, 

Mangold 19 zb (F). Wilson Primitive Reserve, c. 55 km NW of Wauchope, Hale 59237 (US). New England NP., 

Hale 58790, 58794 (US). Mars Rd., Mt. Hyland Nature Reserve, 33 km NW of Dorrigo, Streimann 60628 

(CANB). 15 km W of Dorrigo, on the Armidale Rd., Elix 2332 (CANB). C. 1 km W of Mt. Banda Banda, 

Kantvilas 479/88 (NSW). Mt. Boss SF., 60 km WSW Port Macquarie, Mt. Banda Banda Flora Reserve, at the 

crossing of Banda Rd., and Loop Rd., Wedin 3536 (UPS). Gloucester Tops, Kantvilas 399/88 (NSW). Allyn 

River Forest Park, start of track to Burraga, Kantvilas s.n. (NSW). Blue Mountains NP., Mt. Wilson, Chimney 

Cottage, K. & A. Kalb 20457 (hb. Kalb). Near Blackheath, Blue Mnts., below Bridal Veil Falls, Hale 58679, 

58683 (US). Below Katoomba Falls-trail to Giant Stairway, Katoomba, Hale 58717, 58733 (US). Moreton NP., 

Yurnga Lookout, 35 km NE of Braidwood, Curnow 64 (CANB). SE of Clyde Summit, 37 km from Bateman's 

Bay on main rd. to Braidwood, Hale 69202 (US). Braidwood District, south side of Monga Mt., above 

Mongarlowe River SW of Monga sawmill, 21.07.1968, Weber & Adams s.n. (CANB, COLO, H). Monga 

NP./SF.: 18 km SE Braidwood, along Milo Rd. after turnoff from River Rd., Wedin 3274 (UPS); 27 km SE of 

Braidwood, Mangold 11 k, s (F); Mongarlowe River Area, 4-6 km S of Monga: Elix 11722, 11745, 22679, 

30220, 30241, 30243, Lumbsch 5649 b, Verdon 1317 (CANB); Streimann 15695 (B, CANB). Budawang Range, 

The Vines, near Vine Creek, 12 km from Sassafras, 14 km SE of Nerriga, Verdon 2559 (CANB). 

Chapsa megaphlyctidioides Mangold spec. nov. ined. 

Type: Australia, Queensland, Cape Tribulation, 5 km on Buchanan Creek Rd., Hale 831619 (US-holotype). 

ETYMOLOGY – The epithet refers to the conspicuous, large and distinctly Geaster-like 

ascomata (from gk.: megas =large, great) which distinguishes this species from the otherwise 

similar species C. phlyctidoides. 


Thallus epi- to hypophloedal, (moderately) thin, up to c. 150 µm high, olive to yellowishgreen. 

Surface moderately shiny to waxy, smooth, continuous to slightly verrucose, 

unfissured. True cortex present, continuous, up to c. 50 µm thick, hyaline to pale yellowish, 

consisting of periclinal to rarely irregular hyphae. Algal layer continuous, well developed, 

calcium oxalate crystals variable, rare to lacking in main thallus, abundant near ascomata, 

small to moderately large, clustered. Vegetative propagules not seen. Ascomata conspicuous, 

large, up to c. 2 mm in diam., roundish to slightly irregular, often appearing irregular or 

slightly branched in fused ascomata, distinctly chroodiscoid, erumpent, solitary to fused, 

sometimes regenerating, immersed to slightly raised. Disc usually entirely visible from 

surface, pale flesh-colored to pale grayish, pruinose. Proper exciple not visible from surface 

(inner thalline rim layer may be confused with proper exciple!), thalline rim margin split and 

±lobed, in older ascomata becoming moderately layered, whitish, finely pruinose, ±brownish 

towards the outside, becoming erect to recurved. Exciple fused, thin, hyaline internally, 

yellowish-brown marginally, apically with pale grayish-brown granules, non-amyloid.


Fig. 26. Chapsa magaphlyctidioides: growth habit (A), ascomata (B), ascoma section (C) and 

ascospores (D, E). A.-E.: US-holotype. Bar= A: 2.5 mm; B: 1.25 mm; C: 100 µm; D: 3 µm; E: 4 

µm. 

Hymenium up to c. 70 µm high, non-inspersed, weakly conglutinated, paraphyses slightly 

bent, parallel to slightly interwoven, unbranched, tips unthickened to moderately thickened 

and somewhat irregular, lateral paraphyses present, conspicuous, up to c. 20 µm long. 

Epihymenium moderately thick, hyaline, with grayish granules and sometimes with small 

crystals. Asci 8-spored, tholus (moderately) thin, not visible at maturity. Ascospores (very) 

small, transversely septate, cell walls and endospore thin, halo moderately thick, hyaline, nonamyloid, 

fusiform to clavate, ends subacute to acute, loci angular to slightly roundish, 

predominantly ±rectangular, end cells predominantly 

conical, septae (moderately) thin, 

irregular, 10-18 x 3-5 µm with 4-6(8) loci. Pycnidia 

not seen. 


C-, PD+ orange; containing stictic, constictic 

(majors), hypoconstictic, cryptostictic, α-acetylhypoconstictic 

and hypostictic (traces) acids. 

ECOLOGY AND DISTRIBUTION – Chapsa megaphlyctidioides 

was collected in Australia on tree 

bark in a tropical rainforest at sea level. It is only 

known from the type collection in Northern 

Queensland. 

NOTES – This taxon is characterized by the 

smooth, ±shiny to waxy, corticate thallus, large, 

conspicuous ascomata, small, transversely septate, 


C. megaphlyctidioides.


hyaline, non-amyloid ascospores without thickened parts and the stictic acid chemosydrome. 

It is similar to C. phlyctidioides, which differs by a dull, ecorticate thallus, smaller, less 

distinctly Geaster-like ascomata with less layered margins, and larger (up to 24 µm long, with 

up to 8 loci) ascospores, that are non-halonate and have thickened cell walls and endospore. 


Chapsa minor (Kantvilas & Vezda) Mangold comb. nov. ined. 

Bas.: Chroodiscus minor Kantv. & Vezda, Lichenologist 32: 341 (2000). Type: Australia, Tasmania, 

Weindorfers Forest, Kantvilas 16/88 (HO!-holotype; hb.Vezda-isotype). 


Fig. 28. Chapsa minor: growth habit (A), ascomata (B), asci (C) and ascospores (D). A.-D.: HOholotype. 

Bar= A: 5 mm; B: 0.8 mm; C: 9 µm; D: 5 µm. 

Thallus endo- to hypophloedal, thin to very thin, up to c. 80 µm high, grayish-green. 

Surface dull to slightly shiny, smooth, continuous, unfissured to slightly cracked. Thallus 

covered by a continuous to incontinuous protocortex, sometimes becoming distinctly 

conglutinated, forming an incontinuous true cortex consisting of periclinal hyphae, up to c. 10 

µm high. Algal layer discontinuous and poorly developed, calcium oxalate crystals absent. 

Vegetative propagules not seen. Ascomata conspicuous, (moderately) large, up to c. 1 mm in 

diam., roundish to slightly irregular, apothecioid to chroodiscoid, sessile, solitary to rarely


slightly fused, becoming emergent, depressed-urceolate to cupular. Disc partly to rarely 

entirely visible from surface, pale flesh-colored, epruinose. Proper exciple not visible from 

surface, thalline rim margin characteristic, up to 0.5 µm in diam., radially split, rugged and 

lobed, becoming distinctly layered, off-white, predominantly incurved, only outer layers 

becoming erect to recurved. Exciple fused, hyaline internally to yellowish-brown marginally, 

amyloid in basal parts. Hymenium up to c. 80 µm high, non-inspersed, strongly conglutinated, 

paraphyses slightly bent, parallel to slightly interwoven, unbranched, tips unthickened to 

slightly thickened, lateral paraphyses present, inconspicuous, up to c. 20 µm long. 

Epihymenium thin, hyaline to pale yellowish, without granules, often with small crystals. 

Asci 8-spored, tholus moderately thick, not visible at maturity. Ascospores small, transversely 

septate, cell walls (moderately) thin, endospore moderately thin to slightly thickened, with 

thin halo only in young stages, hyaline, non-amyloid to faintly amyloid ellipsoid to more 

often fusiform to clavate with narrowed-rounded to usually (sub)acute ends, loci roundish to 

angular, subglobose to rectangular or irregular, end cells hemispherical to conical, septae 

thick, regular, 20-28 x 4-6 µm with 6 to 9 loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellowish to brown, C-, PD+ orange; containing stictic, 

hypostictic (majors to minors), constictic (minor to trace), and cryptostictic (traces) acids. 

ECOLOGY AND DISTRIBUTION – Chapsa minor 

was collected in Australia on tree bark in cool 

temperate rainforests in altitudes ranging from 300 

to 1000m. It is a rare species occurring in Tasmania 

and southern Victoria, not yet reported from 

outside Australia. 

NOTES – This taxon is characterized by the thin 

thallus and the bright, apothecioid to indistinctly 

Geaster-like ascomata with layered margins and 

epruinose discs, the moderately small, transversely 

septate, hyaline, non- to faintly amyloid ascospores 

with thin cell walls and thickened endospore and 

septae and the presence of the stictic acid 

chemosyndrome. A similar, and probably closely 

related species is C. lamellifera, which agrees in 

having layered ascomata and containing the stictic 

acid chemosyndrome. It can be readily 

distinguished, however, by larger, muriform 

ascospores. For differences to C. phlyctidioides see 

under this species. 

SPECIMENS EXAMINED – Australia, Victoria: Warburton, Feb. 1902, Wilson s.n. (MEL n. 26169). 

Chapsa niveocarpa Mangold spec. nov. ined. 


C. minor. 

Type: Australia, Queensland, Atherton Tablelands, Tumoulin Rd., 5 km from turnoff to Ravenshoe, Lumbsch 

& Mangold 19151 p (CANB-holotype; BRI-isotype). 

ETYMOLOGY – The epithet refers to the inspersed hymenium and the whitish-pruinose 

surface of the ascomata (from lat.: niveus =snowy and gk. -carpus =fruit).



Fig. 30. Chapsa niveocarpa: ascomata (A, B), ascospore (C) and ascospore detail (D). A.-D.: CANBholotype. 

Bar= A, B: 0.7 mm; C: 20 µm; D: 15 µm. 

Thallus predominantly hypo- to somewhat epiphloedal, (moderately) thin, up to c. 200 µm 

high, pale gray to pale grayish-green. Surface dull, rough to somewhat pruinose, often with 

protuberant substrate structure, continuous, unfissured. Cortex structures predominantly 

absent, thallus rarely covered by a thin, incontinuous protocortex up to 20 µm. Algal layer 

poorly to moderately well developed, predominantly incontinuous, calcium oxalate crystals 

moderately abundant, usually small, clustered to more often scattered. Vegetative propagules 

not seen. Ascomata conspicuous, (moderately) large, up to c. 2 mm in diam., roundish to 

slightly irregular, particularly in fused ascomata, apothecioid to chroodiscoid in older 

ascomata, erumpent, solitary to fused, sometimes regenerating, immersed. Disc partly to 

entirely visible from surface, pale grayish to whitish, strongly pruinose. Proper exciple not 

visble from surface to sometimes visible when becoming partly detached, whitish, thalline 

rim margin irregular, usually coarsely split and lobed, sometimes eroded, slightly to distinctly 

layered, distinctly whitish-pruinose on the insides, becoming erect to ±recurved. Proper 

exciple thin to evanescent, hyaline internally to pale yellowish to pale orange marginally, with 

substrate layers incorporated, apically covered by a usually thick layer of grayish granules, 

non-amyloid. Hymenium up to c. 200 µm high, interspersed by granules (see below), weakly 

conglutinated, paraphyses distinctly adspersed by fine, hyaline granules, straight to slightly 

bent, parallel to slightly interwoven, unbranched, tips distinctly thickened, lateral paraphyses 

present, conspicuous (but often obscured by granule inclusions), up to c. 45 µm long, 

adspersed with fine granules. Epihymenium thick to very thick, hyaline with coarse grayish 

granules. Asci 1- to very rarely 2-spored, tholus thin, not visible at maturity. Ascospores 

(moderately) large, densely eumuriform, cell walls and endospore thin, with thin to


moderately thick, then irregular halo, hyaline, becoming somewhat yellowish to grayish at 

late maturity, non-amyloid, oblong to slightly fusiform, with roundish to narrowed-roundish 

ends, loci small, angular to slightly roundish, with distinct, regular, thin, transverse septae in 

younger ascomata, becoming somewhat irregular in older ascospores, 80-190 x 20-50 µm 

with multiple loci. Pycnidia not seen. 


ECOLOGY AND DISTRIBUTION – Chapsa niveocarpa 

was collected in Australia on tree bark in 


1100 m. It is a rare species occurring to Northern 

Queensland. 

NOTES – The interspersed hymenium, caused by 

the adspersion of the paraphyses with fine granules 

is the most characteristic feature of this species, a 

character that is also occurring in C. leprocarpa 

and C. patens, where granules can be found 

restricted to the lateral paraphyses. Chapsa 

niveocarpa is further characterized by a thin, 

ecorticate thallus, strongly pruinose, whitish 

ascomata, large, densely eumuriform, hyaline, nonamyloid 

ascospores without thickened parts and the 

lack of secondary metabolites. Of those taxa only 

C. leprocarpa occurs in Australia. It can be 

distinguished by a thinner thallus, less strongly 

pruinose ascomata with a lower hymenium (up to 

140 µm high) and smaller ascospores (up to 130 µm long) that more often occur by two per 

ascus. C. patens is morphologically similar, but lacks granular paraphyses and has slightly 

smaller ascospores (up to 160 µm long). It does not occur in Australia. 

SPECIMENS EXAMINED – Australia, Queensland: Mt. Lewis Rd., W of Mossman, Hale 832394 (US). Atherton 

Tablelands: Lake Eacham NP., track around lake, Hale 831188 (US); Palmerston NP., 6 km E of the west 

boundary, Hale 831721 (US); Tumoulin Rd., 5 km from turnoff to Ravenshoe, Mangold 30 p (F); 13 km S of 

Ravenshoe on Tully Falls Rd., Hale 831974 (US). Dawson logging area, WSW of Tully, Hale 832138 (US). 

Chapsa phlyctidioides (Müll.Arg.) Mangold comb. nov. ined. 

Bas.: Ocellularia phlyctidioides Müll.Arg., Hedwigia 32: 130 (1893). Thelotrema phlyctidioides (Müll.Arg.) 

Hale, Mycotaxon 11: 132 (1980). Type: Australia, Queensland, Brisbane, Bailey 354 (G!-holotype). 



C. niveocarpa. 

Thallus epi- to hypophloedal, thin, up to c. 150 µm high, pale gray to greenish-gray. 

Surface dull, smooth to slightly roughened, continuous to verrucose or verruculose, often 

fissured. True cortex absent, thallus covered by an indistinct, incontinuous protocortex up to 

c. 10 µm thick. Algal layer ±continuous but poorly developed, calcium oxalate crystals 

variable, sparse to abundant, small to large, singular or clustered. Vegetative propagules not 

seen. Ascomata conspicuous, predominantly moderately large, up to c. 0.8 mm in diam., 

roundish to slightly irregular, often appearing slightly branched or irregular in fused 

ascomata, apothecioid to indistinctly chroodiscoid, erumpent, solitary to fused, regenerating,


immersed to slightly emergent. Disc partly visible from surface, pale flesh-colored, pruinose, 

in deceased ascomata disc becoming exposed, turning into a whitish-pruinose layer which 

resembles the thalline rim of subsequent ascomata. Proper exciple not visible from surface 

(inner thalline rim layer may be confused with proper exciple!), thalline rim margin structure 

variable throughout ontogeny, in newly developing ascomata split, rugged to lobed, 

±concolorous with thallus or brighter, incurved to erect, in subsequent ascomata generations 

margin layered, splitting and opening irregularly, breaking away in some parts, remaining 

parts finally forming the new inner thalline rim, whitish-pruinose, incurved to erect, slightly 

recurved only in outer layers. Proper exciple fused, thin to evanescent, hyaline internally to 

pale yellowish marginally, distinctly amyloid in marginal parts, amyloidity extending towards 

lateral subhymenial margins. Hymenium up to c. 80 µm high, non-inspersed, in older 

ascomata often with crystal inclusions, moderately conglutinated, paraphyses slightly bent, 

parallel to slightly interwoven, unbranched, tips slightly thickened, somewhat irregular, 

lateral paraphyses present, inconspicuous, up to c. 20 µm long. Epihymenium moderately thin 

to moderately thick, hyaline with yellowish-gray granules and crystals. Asci 8-spored, tholus 

moderately thick, not visible at maturity. Ascospores small, transversely septate, very rarely 

with a single longitudinal septum, cell walls and endospore moderately thick, non-halonate, 

hyaline, non-amyloid to faintly amyloid, ellipsoid to fusiform to clavate, ends roundish to 

subacute, loci roundish to slightly angular, subglobose to more often lentiform or acutelentiform 

with hemispherical to conical end cells, septae thin to moderately thick, regular, 15- 

24 x 5-7µm with 4-8 (x2) loci. Pycnidia not seen. 

Fig. 32. Chapsa phlyctidioides: growth habit (A), ascomata (B) and ascospores (C, D). A., D.: Hale 

831193; B., C.: Elix 18609. Bar= A: 2 mm; B: 1 mm; C: 10 µm; D: 8 µm.


CHEMISTRY – Thallus K+ yellowish to brown, C-, PD+ orange; containing stictic (major), 

constictic, hypostictic (majors to minors), cryptostictic and α-acetylhypoconstictic (traces) 

acids. 


phlyctidioides was collected in Australia on tree 

bark in (sub)tropical rainforests in altitudes ranging 

from 10 to 800 m. It is a moderately common 

species occurring in northern Queensland, the 

Queensland/New South Wales border region and 

on Norfolk Island. Besides Australia the species 

was reported from Dominica, Central America and 

Sri Lanka (Hale, 1981) indicating a pan(sub)tropical 

distribution. 

NOTES – This taxon is characterized by the thin, 

dull, ecorticate thallus, almost closed, regenerating 

ascomata, small, transversely septate, hyaline 

ascospores and the presence of the stictic acid 

chemosydrome. Chapsa minor is similar, 

containing stictic acid, but having raised, regular, 

solitary and epruinose ascomata. Chapsa esslingeri 

(Hale) Mangold comb. nov. ined. 3 from Panama is 

similar, but has distinctly emergent, solitary 

ascomata, an inspersed hymenium and strongly amyloid ascospores that tend to turn slightly 

brownish with age. 

SPECIMENS EXAMINED – Australia, Queensland: S of Noah's Beach on the Cape Tribulation Rd., Hale 

832275 (US). Atherton Tablelands: Malanda Falls, Lumbsch & Mangold 19129 s (F); Curtain Fig Tree S.F.P., 3 

km S of Yungaburra, Hale 831624 (US); Souita Falls, Lumbsch & Mangold 19155 u (F). Babinda Boulders, 

Mangold 39 l, v, x, ze (F). State Forest on Tully Rd., 1 km from jct. with S. Mission Beach Rd., S of Mission 

Beach, Hale 831725 (US). Eungella NP., along Broken River, Lumbsch & Mangold 19100 f (F). Noosa NP., E 

of Noosa Heads, Hafellner 16694 (GZU). Kenilworth SF., SW of Kenilworth, Hale 831193 (US). Lamington 

NP.: Main Border Track out of O'Reillys, Hale 831729, 832269 (US); Python Rock Track, Hale 830863 (US). 

New South Wales: Lions Tourist Rd. near Queensland border, N of Wiangaree, Hale 832712 (US). Mt. Warning 

NP., W of Murwillumbah, Hale 832115 (US). Norfolk Island: Mt. Pitt Reserve, Red Road Track to Mt. Bates, 

Elix 18609 (CANB). 

Chapsa platycarpa (Tuck.) A. Frisch 


C. phlyctidioides. 

Bibl. Lichenol. 92: 113 (2006). Bas.: Thelotrema platycarpum Tuck., Proc. Amer. Acad. Arts Sci. . 5: 406 

(1862). Phaeotrema platycarpum (Tuck.) Zahlbr., Cat. Lich. Univ. II: 609 (1923). Type: Cuba, Wright, Lichenes 

Cubae no. 139 (FH-Tuck.-holotype; BM!-, H-Nyl. 22667-, G-, L-, M-, PC-, UPS-, US!-, W-isotypes). 

Thelotrema platycarpoides Tuck., Proc. Amer. Acad. Arts Sci. . 6: 270 (1864). Phaeotrema platycarpoides 

(Tuck.) Müll. Arg., Flora 69: 311 (1886). Type: Cuba, Wright, Lichenes Cubae no. 157 (FH-Tuck.-lectotype, 

selected by Frisch [2006: 116 - 'piece 6']; BM-, FH-Tuck.-, G-, H-Nyl. 22671-, L-, M!-, PC-, UPS-, US!-, Wisolectotypes). 

Phaeotrema apertum C. W. Dodge, Nova Hedwigia, Beiheft 12: 98 (1964). Type: Uganda, Dummer 4293 

(BM!-holotype; FH-Dodge-isotype). 

[For additional synonymy see Frisch (2006).] 

3 Chapsa esslingeri (Hale) Mangold comb. nov. ined. Bas.: Ocellularia esslingeri Hale, Smithson. Contrib. Bot. 

38: 20 (1978). Thelotrema esslingeri (Hale) Hale, Mycotaxon 11: 131 (1980). Type: Panama, Veraguas, T. L. 

Esslinger 4626 (US!-holotype). [Hale (1978: 20) reports an unidentified PD negative compound in this taxon, 

however, the chemical test of the type material resulted in stictic and constictic acids as major compounds.]



Fig. 34. Chapsa platycarpa: ascomata (A), section of ascoma margin (B) and ascospores (C-E). A., 

E.: Hale 830990; B.: BM-isotype; C., D.: US-isolectotype of T. platycarpoides. Bar= A: 1.25 mm; 

B: 20 µm; C, E: 7 µm; D: 10 µm. 

Thallus hypo- to epiphloedal, (moderately) thin, up to c. 300 µm high, dark olive to olivebrown 

or pale yellowish-brown. Surface dull to wax-like, smooth, continuous to verrucose, 

unfissured. True cortex present, up to 50 µm thick, continuous, yellowish, formed by 

periclinal to irregular hyphae, sometimes with crystal inclusions. Algal layer continuous and 

well developed, calcium oxalate crystals abundant, mostly small, and scattered, sometimes 

clustered. Vegetative propagules not seen. Ascomata conspicuous, (moderately) large, up to c. 

2 mm in diam., roundish to slightly irregular, apothecioid to chroodiscoid in older ascomata, 

erumpent, solitary to fused, often clustered in groups of c. 2-6 ascomata, immersed. Disc 

partly to rarely entirely visible from surface, grayish, distinctly pruinose. Proper exciple 

usually almost entirely visible from surface, forming an inner margin/pore, slightly split to 

ridged, free, irregular, wide to gaping, whitish to off-white, often ±shrunken, predominantly 

incurved to rarely somewhat erect. Thalline rim margin split and coarsely lobed or eroded, 

off-white to whitish on the insides, concolorous with thallus on the outside, erect to recurved. 

Proper exciple free, (moderately) thin, hyaline internally to pale yellowish-brown or brownish 

marginally, non-amyloid. Hymenium up to c. 100 µm high, non-inspersed, moderately 

conglutinated, paraphyses straight to slightly bent, parallel to slightly interwoven, 

unbranched, tips distinctly thickened and somewhat irregular, lateral paraphyses present, 

conspicuous, up to 20 µm long. Epihymenium (moderately) thick, hyaline, with grayish to 

brownish granules and small crystals. Asci 8-spored, tholus moderately thick to moderately 

thin, not visible at maturity. Ascospores (very) small, transversely septate, cell walls and


endospore moderately thick, non-halonate, brown, weakly to moderately amyloid, ellipsoid to 

fusiform with roundish-narrowed to subacute ends, loci roundish to angular, roundishlentiform 

to oblong or irregular, end cells conical, septae (moderately) thick, regular, 10-20 x 

4-6 µm with 4-6(7) loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellow, C-, PD+ orange; containing constictic and stictic 

(majors) and α-acetylconstictic (trace) acids. 


platycarpa was collected in Australia on tree bark 

in (sub)tropical rainforests and monsoon forests in 

low altitudes ranging from sea level to 100 m. It is 

a rare species, known from north-western Northern 

Territory (Frisch, 2006), northern and central 

Queensland. Besides Australia it was reported from 

the Neotropics, Africa, India, Sri Lanka and SE 

Asia (for references see Frisch, 2006) indicating a 

pan(sub)tropical distribution. 

NOTES – This taxon in characterized by a dark, 

corticate thallus, chroodiscoid ascomata with an 

incurved, free proper exciple, small, transversely 

septate, brown, indistinctly amyloid ascospores 

with thickened parts and the stictic acid 

chemosyndrome compounds. The only similar 

Australian species is C. leprieurii, see there for 

differences. Another similar species is Thelotrema 

neei from Panama, which differs by a hypophloedal 

thallus, ascomata with epruinose, reddish-brown discs and smaller (up to 14 µm long, with up 

to 4 loci), non-amyloid ascospores. Although this taxon has chroodiscoid ascomata and 

thereby could be combined to Chapsa, I prefer to keep it tentatively in Thelotrema since it has 

a distinctly free proper exciple and it is uncertain if Chapsa s. str. is characterized by a fused 

to only indistinctly free proper exciple (see also notes of the genus description). 

SPECIMENS EXAMINED – Australia, Queensland: Iron Range NP., 3.5 km from western boundary on track to 

Portland Rds., Hale 830077 (US). Conway Range NP., near Shute Harbour-Airlie Beach, Hale 830990 (US). 

Chapsa pulchra (Müll.Arg.) Mangold comb. nov. ined. 

Bas.: Ocellularia pulchra Müll.Arg., Nuovo Giorn. Bot. Ital. 23: 395 (1891). Type: Australia, Queensland, 

Bailey 583 (G!-holotype; BRI-'Shirley Book', p. 21, n. 34 [BRI-AQ721219]!-isotype). 



C. platycarpa. 

Thallus epi- to predominantly hypophloedal, thin, up to 100 µm high, pale gray to pale 

grayish-green. Surface dull, rarely smooth to usually rough with protuberant substrate surface, 

continuous, unfissured, sometimes appearing fissured due to substrate structure. Cortex 

structures absent. Algal layer continuous to incontinuous, moderately well to poorly 

developed, calcium oxalate crystals sparse to abundant, small to large, scattered or clustered. 

Vegetative propagules not seen. Ascomata variable, conspicuous, moderately large to very 

large, up to c. 2 mm in diam., roundish to irregular, particularly in fused ascomata, apo- to


chroodiscoid in older stages, erumpent, solitary to fused, often clustered in groups of few to 

many ascomata, then usually smaller, immersed. Disc partly to entirely visible from surface, 

pale flesh-colored to grayish, usually with distinct pruina and ±glittering. Proper exciple not 

visible from surface, thalline rim margin (moderately) thick, split to somewhat lacerate, lobed 

to eroded, rarely slightly layered, ±distinctly pruinose, whitish, becoming erect to ±recurved. 

Exciple fused, thin to evanescent, hyaline internally to yellowish or pale orange marginally, 

often with incorporated substrate particles, apically usually covered in grayish to brownishgray 

granules, non-amyloid. Hymenium up to c. 130 µm high, non-inspersed, highly 

conglutinated, paraphyses straight, parallel to slightly interwoven, unbranched, tips slightly 

thickened and somewhat irregular, lateral paraphyses present, conspicuous, up to c. 30 µm 

long. Epihymenium (moderately) thick, hyaline, with coarse grayish to brownish-gray 

granules and crystals. Asci 6 to 8-spored, tholus (moderately) thick, thin when mature. 

Ascospores moderately small to moderately large, transversely septate, fragile, often breaking 

apart in microscopic slide, cell walls (moderately) thin, endospore thin to very thin, in earlier 

stages with indistinct but thick to very thick, often irregular halo, hyaline, non-amyloid, 

bacillar to bacillar-fusiform, often ±bent, loci roundish to slightly angular, predominantly 

roundish-rectangular to depressed-rectangular with usually hemispherical end cells, septae 

thin, distinctly regular, 30-60(70) x 5-8 µm with 12-22(24) loci. Pycnidia not seen. 


Fig. 36. Chapsa pulchra: growth habit (A, C), ascomata (B), section of ascoma margin (D) and 

ascospore (E). A.: Mangold 23 u; B., E.: Lumbsch & Mangold 19129 t; C.: Lumbsch & Mangold 

19151 za; D.: G-holotype. Bar= A: 1.75 mm; B: 1 mm; C: 2 mm; D: 30 µm; E: 10 µm.


ECOLOGY AND DISTRIBUTION – Chapsa pulchra 

was collected in Australia on tree bark of several 

trees in predominantly open sites of (sub)tropical 

wet sclerophyll forests and rainforests in altitudes 

ranging from 10 to 1130 m. It is a common species, 

known from Queensland and the Queensland-New 

South Wales border region. So far not reported 

from outside Australia. 

NOTES – It is a quite uniform species and it can 

even be identified in the field by its conspicuous 

whitish ascomata and the endophloedal, bright 

thallus. Only the ascomata are somewhat variable 

regarding their size and arrangement. Older, 

solitary ascomata are usually very large, in groups 

of clustered ascomata they can be found to be 

considerably smaller (see fig. 36, B and C). It is 

further characterized by the large, transversely 

septate, hyaline, non-amyloid, thin-walled, 

±bacillar and fragile ascospores and the lack of 

secondary compounds. In Australia, the closest taxa is C. indica, for differences see there. 

Morphologically similar is C. colobicus, which can be readily distinguished by mono- to 

bisporic asci with densely eumuriform ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Fitzroy Island on Great Barrier Reef, 25 km E of Cairns, A. 

& M. Aptroot 22334/I (ABL). Atherton Tablelands: Lake Tinaroo, Downfall Creek Camping Area, Lumbsch & 

Mangold 19123 j, 19125 h (F); Dunbulla Forest Drive, near Lake Durango parking lot, Mangold 37 j (F); W side 

of Lake Eacham NP., 3 km in from W boundary on rd. from Yungaburra, Hale 832154 (US); Lake Eacham NP., 

Mangold 29 ag (F); Plath Rd. logging head, 9 km W of Plath Rd., off Kennedy Hwy., Herberton Range, S of 

Atherton, Hale 832165 (US); Malanda Falls, Lumbsch & Mangold 19129 t (F); Tumoulin Rd., 5 km from turnoff 

to Ravenshoe, Lumbsch & Mangold 19133 d, I, 19148 a, d, 19151e, k, za (F). Dawson logging area, 24 km S of 

Koombooloomba turnoff, WSW of Tully, Hale 830695 (US). Blencoe Creek, Cardwell Range, 48 km NW of 

Cardwell, Elix & Streimann 20144 (CANB). Dawes Range, Kroombit SF., 53 km E of Biloela, Elix 34751 

(CANB). Kalpowar Forest Drive, 40 km NE of Monto, SW of Gladstone, Hale 831272, 831666, 832515 (US). 

Imbil SF., 6 km NW of Imbil, Rogers 2525 (BRI). Imbil Forest Drive, between stop #4 and #5 on rd. from 

Borumba Dam to Imbil, SW of Glympie, Hale 831366 (US). 31 km S of Gympie, 5 km W of Imbil, Tibell 

12606, 12618 (UPS). Booloumba Creek SF., SW of Kenilworth, Hale 831715 (US). 6 km N of Jimna, Tibell 

12783 (UPS). Sunshine Coast Hinterland, N side of Lake Baroon Pocket, W of Montville, Lumbsch & Mangold 

19082 e (F). Mt. Mee SF.: near Mt. Mee, N of Brisbane, Hale 58561, 58633 (US); 6 km NW of Forestry Office, 

NW of Mt. Mee, Hale 830286, 832116, 832389 (US). Goodna (E of Ipswich, nr. Brisbane): Wilson "1457" (as T. 

megalophthalmum) (NSW-539388); Wilson s.n., Aug. 1889 (NSW-539347). Cunninghams Gap, N of 

Cunningham Hwy., A. & K. Kalb 34260 (hb. Kalb). New South Wales: Tweed Range, Mebbin NP., 25 km SW 

of Murwillumbah, Mangold 21 r, s (F). Nightcap Forest Drive: 1 km W of Minyon Falls, N of Lismore Hale 

832364 (US); Big Shrub Flora Reserve, W of Mullumbimby, Hale 831698, 832040 (US). 'Woolongbar' 

(=?Wollongbar, nr. Lismore), Wilson "1457" (as T. megalophthalmum), Jul.1984 (NSW-539369). Iluka Nature 

Reserve, 50 km NE of Grafton, Mangold 23 u (F). 

Chapsa subpatens (Hale) Mangold comb. nov. ined. 

Bas.: Thelotrema subpatens Hale, Bull. Br. Mus. nat. Hist. (Bot.) 8: 269 (1981). Type: Sri Lanka, Southern 

Province, Galle, Hale 46208 (US-holotype, BM!-isotype). 



C. pulchra.


Fig. 38. Chapsa subpatens: growth habit (A), ascoma (B), ascospores (C, D) and ascospore with 

amyloid reaction (E). A.: BM-isotype; B.-E.: Hale 68794. Bar= A: 2.5 mm; B: 0.4 mm; C: 3 µm; 

D, E: 3.5 µm. 

Thallus predominantly hypophloedal, very thin, up to c. 50 µm high, (pale) grayish. 

Surface dull to slightly shiny, smooth, continuous, predominantly unfissured, sometimes 

appearing fissured due to substrate structure. Cortex structures predominantly absent, rarely 

thallus covered by a thin, incontinuous protocortex up to c. 10 µm thick. Algal layer poorly 

developed, incontinuous, calcium oxalate crystals absent. Vegetative propagules not seen. 

Ascomata inconspicuous, large, up to c. 1.5 mm in diam., roundish to slightly irregular, 

particularly in fused ascomata, apothecioid to chroodiscoid in older ascomata, erumpent, 

solitary to fused, immersed. Disc partly to rarely entirely visible from surface, (pale) 

brownish to grayish-brown, becoming slightly pruinose with age. Proper exciple often visible 

from surface, usually appearing free, but always fused with substrate/thalline rim layers, 

whitish, thalline rim margin irregular, rugged and lobed, lobes large and thin, sometimes 

eroded, often appearing layered due to exfoliating substrate, concolorous with thallus or 

brownish due to protuberant substrate layers, becoming erect to recurved in older stages. 

Exciple (moderately) thick, hyaline internally to pale yellowish marginally, often with 

substrate layers incorporated, apically usually covered by grayish granules, often slightly 

amyloid (reddish) to distinctly amyloid at the base (purple). Hymenium up to c. 80 µm high, 

non-inspersed, weakly conglutinated, paraphyses straight to slightly bent, parallel,


unbranched, tips unthickened to slightly thickened, lateral paraphyses present, inconspicuous 

in younger ascomata, up to c. 20 µm long, not clearly separated from exciple. Epihymenium 

moderately thick, brownish, with fine to coarse grayish-brown granules. Asci 6 to 8-spored, 

tholus moderately thin, thinning or not visible at maturity. Ascospores small, transversely 

septate, cell walls and endospore (moderately) thick, often with distinctly crenate surface, 

non-halonate, hyaline, distinctly to strongly amyloid, predominantly ellipsoid to oblongfusiform, 

rarely fusiform, with roundish to narrowed-roundish ends, loci roundish to slightly 

angular, predominantly subglobose to oblong, end cells hemispherical to conical, septae 

(moderately) thick, regular, 20-30 x 6-7 µm with 7-10 loci. Pycnidia not seen. 


ECOLOGY AND DISTRIBUTION – Chapsa subpatens 

is known in Australia from one collection on 

tree bark in a cool-temperate Nothofagus forest at 

800 m altitude. It is an extremely rare species in 

Australia, only known from Tasmania. This is the 

first report for Australia, so far it was only reported 

from Sri Lanka. 

NOTES – Chapsa subpatens is an inconspicuous 

species, mainly characterized by the very thin, 

ecorticate thallus, the brownish discs, the small, 

transversely septate, hyaline, amyloid ascospores 

with thickened parts and the lack of lichen 

substances. In the type collection (besides the 

present material from Tasmania the only existing 

collection of this species), the ascomata are small 

and almost apothecioid, somewhat resembling a 

member of Thelotrema. The Tasmanian specimen 

however, is probably fully mature or better 

developed and has large, Geaster-like ascomata. 

The only known similar Chapsa in Australia is C. tibellii from northern Queensland, which 

can be readily distinguished by the thicker, corticate thallus and the larger (up to 45 µm long), 

fusiform ascospores. Further, C. crispata from Brazil is the most closely affiliated species, it 

also differs by a distinctly epiphloedal thallus and has thick margins which are, as well as the 

disc, distinctly whitish-pruinose. 

SPECIMENS EXAMINED – Australia, Tasmania: Murchinson Hwy., 57 km S of Bass Hwy., S of Hellyer 

Gorge, Hale 68794 (US). 

Chapsa tibellii Mangold spec. nov. ined. 

Type: Australia, Queensland, Atherton Tablelands, Mt. Haig, 22 km NE of Atherton, Tibell 15314 (UPSholotype). 

ETYMOLOGY – This species in named after the Swedish lichenologist Leif Tibell who 

collected the type specimen. 



C. subpatens.


Fig. 40. Chapsa tibellii: ascomata (A), ascospore (B) and ascospore with amyloid reaction (C). A.-C.: 

UPS-holotype. Bar= A: 1.5 mm; B: 4 µm; C: 5 µm. 

Thallus epi- to somewhat hypophloedal, thin to usually (moderately) thick, up to c. 400 µm 

high, pale yellowish-gray to (pale) greenish-gray. Surface shiny, smooth, predominantly 

distinctly verrucose to verruculose, unfissured, sometimes coarsely cracked. True cortex 

present, thick, up to c. 50 µm thick, continuous, consisting of periclinal to irregular hyphae. 

Algal layer continuous and well developed, calcium oxalate crystals usually abundant, large 

to more rarely small, clustered. Vegetative propagules not seen. Ascomata very conspicuous, 

(very) large, up to c. 5 mm in diam., roundish to irregular, in fused ascomata sometimes 

slightly branched, chroodiscoid, erumpent, solitary to sometimes fused, immersed to 

somewhat emergent, sometimes with a ±constricted base. Disc predominantly entirely visible 

from surface, whitish-gray to gray, distinctly pruinose, with fine pruina. Inner proper exciple 

not visible from surface to sometimes visible, in particular in older ascomata with recurved 

thalline margins, predominantly fused, sometimes partly becoming slightly detached, whitish, 

thalline rim margin thick, split to rugged or coarsely lobed, becoming erect to recurved, in 

older ascomata sometimes eroded, often slightly pruinose internally, ±whitish or brighter than 

thallus, concolorous with thallus marginally. Exciple fused to apically slightly free, 

moderately thin to moderately thick, pale yellowish internally to yellowish-brown or (dark- 

)brown marginally, apically often dark brown and covered with grayish granules, sometimes 

slightly amyloid at the base. Hymenium up to c. 120 µm high, non-inspersed, moderately 

conglutinated, paraphyses ±straight, parallel, with unthickened to slightly thickened, regular 

to slightly irregular tips, lateral paraphyses present, conspicuous, up to c. 25 µm long, 

subhymenium conspicuous, dark-brown to slightly carbonized. Epihymenium moderately 

thick, hyaline, with ±coarse grayish granules. Asci 8-spored, tholus thick, thin when mature. 

Ascospores typical, (moderately) small, transversely septate, cell walls and endospore thick, 

non-halonate, hyaline, distinctly amyloid in mature ascospores, rarely ellipsoid to 

predominantly fusiform with narrowed-roundish to acute ends, loci roundish to somewhat 

angular, predominantly lentiform to irregular-lentiform with predominantly cone-shaped end-


cells, septae (moderately) thick, often becoming irregular, 25-45 x 7-10 µm with 8-12(14) 



ECOLOGY AND DISTRIBUTION – Chapsa tibellii 

was collected in Australia on tree bark in tropical 

rainforests and a wet sclerophyll forest in elevations 

ranging from 800 - 1200 m. It is a rare species 

only known from the Atherton Tablelands region in 

Northern Queensland. 

NOTES – It is a very characteristic species with a 

shiny, corticate, usually thick, verrucose to 

verruculose thallus with abundant crystal inclusions, 

conspicuous, large, distinctly chroodiscoid 

ascomata with bright-gray, pruinose discs 

and thick thallus margins, a dark subhymenium and 

moderately large, hyaline, transversely septate, 

amyloid, fusiform ascospores with thickened cell 

walls. Chapsa dilatata from the Neotropics 4 is a 

similar species that differs in chemistry (stictic acid 

chemosydrome) and in having smaller ascospores 

(up to 33 µm long, with up to 10 loci). Two species 

from Brazil are similar, both lacking secondary 


C. tibellii. 

compounds and having ±similar sized, hyaline, transversely septate ascospores, C. crispata 5 

and C. elabens 6. Chapsa crispata has smaller ascospores (up to 25 µm long, up to 8 loci) and 

smaller, less distinctly chroodiscoid ascomata with a partly free exciple and a thinner thallus. 

C. elabens can by readily distinguished by the ecorticate, thin thallus, the inspersed 

hymenium and the larger ascospores (up to 50 µm long, up to 20 loci). 

SPECIMENS EXAMINED – Australia, Queensland, Atherton Tablelands: Davies Creek Rd. 17 km S of Kennedy 

Hwy., S of Davies Creek Falls NP., E of Mareeba, Hale 830916 (US); Plath Rd. logging head, 9 km W of Plath 

Rd., off Kennedy Hwy, Herberton Range, S of Atherton, Hale 832395, 832679 (US); 10 km S of Ravenshoe on 

Tully Falls Rd., Hale 831443 (US); Culpa logging area, 13 km from Koombooloomba rd. turnoff, SE of Tully 

Falls, Hale 832705 (US); Dawson logging area, State Forest Reserve 605, 24 km S of Koombooloomba turnoff, 

WSW of Tully, Hale 830881, 831654, 832392 (US). 

4 Hale (1981) reports this taxa from Sri Lanka. However, Hale's collection found in US has large, densely 

eumuriform, non-amyloid ascospores and is an undetermined species of Chapsa close to C. recurva/C. 

zahlbruckneri (both types not seen). 

5 Chapsa crispata (Müll. Arg.) Mangold comb. nov. ined. Bas.: Ocellularia crispata Müll. Arg., Journ. Linn. 

Soc. London, Bot., 30: 452 (1895). Type: Brazil, Rio de Janeiro, (ex hb. Kew 1894), Portella s.n. (G!-lectotype, 

here selected). 

6 Chapsa elabens (Müll. Arg.) Mangold comb. nov. ined. Bas.: Ocellularia elabens Müll. Arg., J. Linn. Soc. 

London, Bot. 30: 452 (1895). Type: Brazil, (ex hb. Kew 1894), s.c. (G!-lectotype, selected by Hale [1972, in 

hb.]).


2. 9. 2. Chroodiscus (Müll.Arg.) Müll.Arg., Lichenes epiphylli novi: 18 (1890). Type species: 

Platygrapha coccinea Leight. [= C. coccineus (Leight.) Müll.Arg.]. Type: Brazil, Spruce 

443 (BM-lectotype). 

Ocellularia sect. Chroodiscus Müll.Arg., Flora 66: 352 (1883a). Phyllophthalmaria sect. Chroodiscus (Müll. 

Arg.) Zahlbr., in Engler-Prantl, Die natürlichen Pflanzenfamilien I, 1: 120 (1905). 

Phyllobrassia Vain., Ann. Acad. Sci. Fenn., ser. A 15(6): 173 (1921a). Type species: Platygrapha mirifica 

Kremp. [= C. mirificus (Kremp.) R. Sant.]. Type: Borneo, Beccari 229 b (M-holotype). 

THALLUS – Crustose, foliicolous, thin, epiphyllous, up to c. 40-50 µm high, pale greenish 

to pale grayish-green. Surface dull to shiny, smooth, continuous to slightly verruculose, 

unfissured. Prothallus absent. True cortex absent, covered by a thin, discontinuous 

protocortex up to c. 10 µm thick. Algal layer well developed, continuous, crystals abundant, 

small to moderately large, forming a ±continuous layer beneath the algal layer. Distinct 

medulla layer absent. Vegetative propagules not known from the Australian taxa, discoid 

isidia known from C. mirificus. 

ASCOMATA – Conspicuous, moderately small to moderately large, up to c. 300-600 µm in 

diam., roundish to rarely slightly elongated or irregular in fused ascomata, chroodiscoid from 

early stages on, non-regenerating, solitary to fused, immersed to slightly raised. Disc visible 

from surface, bright orange to reddish-orange or (pale) grayish to (pale) grayish-brown 

epruinose or pruinose. Proper exciple not visible from surface, thalline rim margin moderately 

thin to moderately thick, rarely entire, otherwise slightly to distinctly split, ±distinctly lobed, 

±eroded, particularly in older stages, slightly incurved to more often erect to distinctly 

recurved. Proper exciple fused, thin to evanescent or moderately thick, non-amyloid. 

Subhymenium indistinct, evanescent, hyaline to slightly grayish. Hymenium non-amyloid, 

discoid, up to c. 50-60 µm high, non-inspersed, clear, moderately to distinctly conglutinated, 

paraphyses thin, straight, parallel to slightly interwoven, unbranched, tips not thickened to 

slightly thickened, lateral paraphyses absent. True columella or columellar structures absent. 

Epihymenium thin to moderately thick, hyaline or pale orange to rust-red, granulose. 

Asci 8-spored, tholus moderately thin to thin or moderately thick, non-amyloid, clavate. 

Ascospores uni- to biseriate, small, 7-15 x 2-4 µm, transversely septate, cell walls moderately 

thin, smooth, non-halonate, hyaline, non-amyloid, ellipsoid to fusiform, straight, with 2-6 

loci, end cells hemispherical to more often conical, loci large, ±angular, cuboid, septae 

moderately thin to thin, regular to irregular. 

PYCNIDIA – Not known. 

CHEMISTRY – Secondary compounds present, containing the stictic acid chemosyndrome 

and unknown anthraquinones. 

ECOLOGY AND DISTRIBUTION – The Chroodiscus species in Australia occur on living 

leaves in altitudes ranging between 20 and 800 m. The distribution is restricted to tropical 

rainforests of the eastern coast of northern Queensland. Both taxa occurring in Australia are 

pantropical. 

NOTES – Besides Hales’ inclusion in Thelotrema (1981), this comparably well-known and 

well separated taxonomic group of Thelotrematacae has been treated as a distinct genus ever 

since Müller's introduction as an independent genus in 1890. Several authors recently (Kalb, 

1991; Kantvilas & Vezda, 2000; Staiger, 2002; Vezda, 1992) included corticolous and 

temperate foliicolous taxa with lateral paraphyses which are now grouped in Acanthotrema, 

Chapsa and Topeliopsis.


The genus is readily characterized by small, foliicolous thalli, distinctly chroodiscoid 

ascomata without lateral paraphyses, small, hyaline, non-amyloid ascospores and a 

distribution strictly restricted to tropical rainforests. Similar genera include Acanthotrema, 

Chapsa and Reimnitzia, all well distinguished by a different ecology, larger thalli and (except 

the latter, see below) lateral paraphyses. The presence of lateral paraphyses in Reimnitzia is 

uncertain (see also there), however, this monotypic genus is further distinguished from 

Chroodiscus by a thick and bulging thallus, slightly branched hymenia and brown ascospores. 

The report of C. mirificus for Australia (Lumbsch & Vezda, 1990) was based on a 

misidentification of C. parvisporus. 


Chroodiscus australiensis Vezda & Lumbsch 

Nova Hedwigia 50: 246 (1990). Type: Australia, Queensland, Atherton Tableland, Souita waterfalls, 

Lumbsch 5437/17 ei, (F!-holotype, hb. Vezda-isotype). 


Fig. 42. Chroodiscus australiensis: growth habit (A), ascomata (B) and asci and ascospores (C, D). 

A.-D.: F-holotype. Bar= A: 1 mm; B: 275 µm; C: 7.5 µm; D: 5 µm. 

Thallus epiphyllous, thin, up to c. 40 µm high, pale greenish. Surface dull to shiny, 

smooth, continuous, unfissured. True cortex absent, covered by a thin, incontinuous 

protocortex up to c. 10 µm thick. Algal layer well developed, continuous, crystals abundant,


small to moderately large, forming a ±continuous layer beneath the algal layer. Vegetative 

propagules not seen. Ascomata conspicuous, moderately small, up to c. 300 µm in diam., 

roundish, chroodiscoid, solitary to marginally slightly fused, immersed to slightly raised. Disc 

predominantly entirely visible from surface, bright orange to reddish-orange, epruinose. 

Proper exciple not visible from surface, thalline rim margin moderately thin, entire to slightly 

split, ±distinctly lobed only in younger ascomata, in older stages becoming eroded, 

predominantly pale orange, slightly incurved to erect to more rarely recurved. Proper exciple 

fused, thin to evanescent, pale orange, non-amyloid. Hymenium up to c. 50 µm high, noninspersed, 

pale orange, moderately conglutinated, paraphyses straight, parallel to slightly 

interwoven, unbranched, tips unthickened to slightly thickened, lateral paraphyses absent. 

Epihymenium moderately thick, pale orange to rust-red, with rust-red, moderately small 

crystals. Asci 8-spored, tholus (moderately) thin, not visible at maturity. Ascospores very 

small, transversely septate, cell walls moderately thin, endospore lacking, non-halonate, 

hyaline, non-amyloid, ellipsoid to more often fusiform with narrowed roundish to subacute 

ends, loci predominantly conical, in 3-locular ascospores internal locus angular, septae 

(moderately) thin, 7-10 x 2-3 µm with 2(3) loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellowish to brown, C-, PD+ orange; containing stictic and 

hypostictic (majors) acids, ascomata K+ purplish, containing an unknown orange to rust-red 

anthraquinone. 

ECOLOGY AND DISTRIBUTION – Chroodiscus 

australiensis was collected in Australia on living 

leaves in tropical rainforests in altitudes ranging 

between 50 and 700 m. It is a rare species occurring 

in northern Queensland. Besides Australia it was 

reported from the Neotropics, Africa and South- 

East Asia (for references see Frisch, 2006), 


NOTES – This taxa is a conspicuous foliicolous 

lichen that is readily characterized by the reddishorange 

ascomata and the 2(3)-celled ascospores. 

For differences to C. parvisporus, the only other 

member of Chroodiscus in Australia, see under that 

species. On the world level the pantropical 

C. coccineus is morphologically similar, it can be 

distinguished by a distinctly verruculose thallus 

surface and larger ascomata (up to c. 500 µm in 

diam.) and ascospores (up to 12 µm long, with up 

to 4 loci). 

SPECIMENS EXAMINED – Australia, Queensland: Daintree NP., W of Mossman, A. & K. Kalb 25376 (F). 

Atherton Tablelands, Souita waterfalls, Lumbsch 5437 zc, zg, ep, zh (F). Bellenden Ker NP., Boulders area, 

Aptroot 22443, 22458 (B), 22431 (ABL). Palmerston NP., near Tchupala Falls, Aptroot 22580 (ABL). Daintree 

NP., W of Mossman, A. & K. Kalb 25376 (F). 

Chroodiscus parvisporus Kalb & Lücking 


C. australiensis. 

Stapfia 80: 271 (2002). Type: Australia, Queensland, surroundings of Daintree Village, Daintree River, A. & 

K. Kalb s.n. (hb. Kalb-holotype, not seen).



Fig. 44. Chroodiscus parvisporus: growth habit (A), ascomata (B), asci (C) and ascospores (D). A., 

B.: A. & K. Kalb s.n.; C., D.: Lumbsch & Mangold 19126 a. Bar= A: 0.8 mm; B: 250 µm; C, D: 5 

µm. 

Thallus epiphyllous, thin, up to c. 50 µm high, pale grayish-green. Surface dull to slightly 

shiny, smooth, continuous to slightly verruculose, unfissured. True cortex absent, covered by 

a thin, incontinuous protocortex up to 10 µm thick. Algal layer well developed, continuous, 

crystals abundant, small, forming a ±continuous layer beneath the algal layer. Vegetative 

propagules not seen. Ascomata conspicuous, moderately large, up to c. 600 µm in diam., 

predominantly roundish to more rarely slightly elongated, irregular in fused ascomata, 

chroodiscoid, solitary to fused, usually slightly raised. Disc predominantly entirely visible 

from surface, (pale) grayish to (pale) grayish-brown, epruinose to slightly pruinose. Proper 

exciple not visible from surface, thalline rim margin thick, split and ±distinctly lobed, often 

somewhat eroded, off-white, erect to predominantly recurved. Proper exciple fused, 

moderately thick, pale brownish to pale grayish-brown, non-amyloid. Hymenium up to c. 60 

µm high, non-inspersed, distinctly conglutinated, paraphyses straight, parallel to slightly 

interwoven, unbranched, tips unthickened to slightly thickened, lateral paraphyses absent. 

Epihymenium thin to indistinct, hyaline, with a few fine ±colorless granules and small 

crystals. Asci 8-spored, tholus moderately thick throughout development, lateral walls 

moderately thick, thin when mature. Ascospores very small, transversely septate, cell walls 

moderately thin, endospore lacking, non-halonate, hyaline, non-amyloid, oblong to ellipsoid 

with roundish to slightly narrowed roundish ends, loci predominantly angular with conical to 

hemispherical end cells, septae (moderately) thin, irregular, 10-15 x 3-4 µm with 3-6 loci. 

Pycnidia not seen.



constictic (majors), hypoconstictic, hypostictic (minors) and α-acetylconstictic (trace) acids. 

ECOLOGY AND DISTRIBUTION – Chroodiscus 

parvisporus was collected in Australia on living 

leaves in tropical rainforests in altitudes ranging 

between 20 and 800 m. It is a rare species occurring 

in northern Queensland. Besides Australia it was 

reported from the Neotropics and Indomalaysia 

(Lücking & Grube, 2002), indi-cating a pantropical 

distribution. 


grayish to brownish discs, transversely septate, 

relatively large ascospores and the stictic acid 

chemosydrome compounds. Chroodiscus australiensis 

differs in smaller ascospores (up to 10 

µm with up to 3 loci) and reddish, K+ purple discs, 

containing an unknown anthraquinone. 

SPECIMENS EXAMINED – Australia, Queensland: 

Surroundings of Daintree Village, A. & K. Kalb s.n. (F). 

Atherton Tablelands: Curtain Fig Tree, Lumbsch 5427/8-zh 


C. parvisporus. 

(F); Souita Falls, Lumbsch 5437/10-zii, 5437/17-eh (F); Lake Euranoo, Lumbsch & Mangold 19126 a (F). 

2. 9. 3. Fibrillithecis Frisch, Bibl. Lichenol. 92: 135 (2006). Type species: Thelotrema 

vernicosum Zahlbr. [=F. halei (Tuck. & Mont.) Mangold comb. nov. ined.]. Type: U.S.A., 

Hawaii, Oahu, Rock 101 (W-lectotype). 

Thelotrema sect. Tremotyliopsis Zahlbr., Denkschr. Kaiserl. Akad. Wiss. Wien, math.-naturw. Klasse 83: 120 

(1909). Type species: Thelotrema insigne Zahlbr., Denkschr. Kaiserl. Akad. Wiss. Wien, math.-naturw. Klasse 

83: 120 (1909). Type: Brazil, São Paulo. Wettstein & Schiffner s.n. (W-holotype). 

Fibrillithecis halei is the only known species in Australia, for a description see there. 

NOTES – This recently described new genus was introduced to accommodate three taxa (F. 

insignis, F. platyspora, F. vernicosa) predominantly characterized by a distinctly fibrous 

proper exciple. Only two species are accepted here, viz. F. halei and F. insignis which are 

distinguished by the occurrence of isidia. In F. halei isidia are absent, in F. insignis, which is 

not known from Australia, distinct cylindrical, branched isidia are present. Fibrillithecis 

platyspora and F. vernicosa are considered synonymous to F. halei, see notes in the 

description of this species. 

The taxonomic uncertainties in this group are reflected in the combination of Fibrillithecis 

taxa in Hale’s exciple structure-based classification (1980, 1981). The two species where 

either treated as Thelotrema (e.g. as T. piluliferum or T. platysporum [=T. halei]) based on the 

interpretation of the apical excipular fibrils as lateral paraphyses, or Myriotrema (as M. 

insigne [=F. insignis]) due to a stronger emphasis of characters as the perithecioid ascomata 

and the fused proper exciple. Myriotrema is the most similar genus to Fibrillithecis, which is 

also supported by molecular data (Frisch & al., 2006). The two genera are distinguished by 

the structure of the apical proper exciple, in Myriotrema distinct fibrils are absent. Frisch 

(2006: 137) further gives a different structure of the cortex as a separating character.


Species description: 

Fibrillithecis halei (Tuck. & Mont.) Mangold comb. nov. ined. 

Bas.: Porina halei Tuck. & Mont., Annal. Scienc. Nat. Bot. 4(8): 295 (1857). Thelotrema halei (Tuck. & 

Mont.) Nyl., Annal. Scienc. Nat. Bot. 4(11): 221 (1859). Type: Venezuela, Dec. 1838, Fendler s.n. (FH-Tuck.!lectotype, 

selected by Hale [1972 in hb.]; H-Nyl.-isolectotype). 

Thelotrema argenteum Müll.Arg., Hedwigia 30: 50 (1891). Type: Australia, Queensland, Bellenden Ker, 

Bailey 547 (G-holotype; BM!-, BRI!-, C!-, NSW!-, NY!-, TNS!-, WIS!-isotypes). 

Thelotrema pachystomum ssp. piluliferum Tuck., Proc. Amer. Acad. Arts Sci. 7: 227 (1866). Type: U.S.A., 

Hawaii, Oahu, Waiahu Mtns., Mann s.n. (FH-Tuck.!-lectotype, re-selected by Frisch [2006: 140]; G-, Wisolectotypes). 

Thelotrema vernicosum Zahlbr., Ann. Mycol. 10: 370 (1912). Fibrillithecis vernicosa (Zahlbr.) Frisch, Bibl. 

Lichenol. 92: 140 (2006). Type: U.S.A., Hawaii, Oahu, Koolau Mnts., Rock 101 (W-lectotype, selected by Hale 

[1981: 264]; FH!-isolectotype). 

Thelotrema platysporum Harm., Bull. Soc. Sci. Nancy, 3(13): 41 (1912). Fibrillithecis platyspora (Harm.) 

Frisch, Bibl. Lichenol. 92: 137 (2006). Type: ?New Caledonia/Australia ("pro maxima parte in Nova Caledonia, 

pro minima vero in Australia collecti"), Pionniero 38 (DUKE-lectotype, selected by Hale [1981: 266]; FH!-, Wisolectotypes). 

Thelotrema gibbosum H. Magn., in Magnusson & Zahlbruckner, Ark. Bot. 31 A(1): 53 (1943). Type: 

Hawaii, W. Maui, Haelaau, Selling 5836 (UPS-lectotype, selected by Hale [1981: 264]; S-isolectotype). 

Thelotrema diminitum Hale, Phytologia 27: 494 (1974). Type: Sarawak, Bako National Park, Hale 30536 

(US!-holotype). 


Thallus corticolous to rarely muscicolous, epi- to hyposubstratic, moderately thin to 

moderately thick, up to c. 400 µm high, pale greenish- to yellowish-gray, dark olive-gray or 

off-white. Surface ±shiny, smooth to rarely partly somewhat pruinose, predominantly 

±verrucose, usually fissured to more rarely somewhat areolate. True cortex present, 

continuous to sometimes discontinuous, up to c. 30 µm thick, consisting of irregular to 

periclinal hyphae. Algal layer ±well developed, continuous, with ±abundant, small to large 

calcium oxalate crystals, in (very) large clusters. Vegetative propagules not seen. Ascomata 

variable, usually conspicuous, (moderately) large, up to c. 1(1.5) mm in diam., roundish, 

somewhat irregular in fused ascomata, predominantly perithecioid to rarely apothecioid, 

solitary to fused, rarely somewhat clustered in groups, immersed to strongly emergent, then 

(flattened-)cylindrical to (flattened-)urceolate with continuous surface. Disc usually not 

visible from surface, in open to gaping ascomata partly visible, pale flesh-colored, with 

whitish pruina. Pores (very) small and opening at late maturity, sometimes wide to rarely 

gaping, up to c. 200 µm in diameter, usually not larger than c. 80 µm in diam., roundish to 

slightly irregular, pore margin rarely entire to fibrous and formed by the apical proper exciple, 

then usually ±sunken, fused proper exciple often becoming apically visible from surface due 

to evanescent thalline rim margin, concolorous with thalline rim to somewhat grayish or 

brownish, in some specimen proper exciple becomes partly to entirely detached, then visible 

from surface in its main upper parts, with whitish-pruinose surface. Thalline rim margin 

moderately thin to thick, roundish, entire to often ±eroded, small to moderately wide to rarely 

wide or gaping, incurved, predominantly concolorous with thallus. Proper exciple 

conspicuous, predominantly fused, rarely becoming ±free, thick to very thick, forming lateral 

paraphyses-like structures by apically distinctly radiating hyphae, hyaline internally to 

yellowish-brown or orange-reddish marginally, apically sometimes grayish-brown to darkbrown, 

non-amyloid. Hymenium up to c. 180 µm high, non-inspersed, highly conglutinated, 

paraphyses unbranched, parallel, ±bent, irregular and often distinctly multicellular, tips 

thickened, lateral paraphyses and columellar structures lacking. Epihymenium thin, hyaline, 

with fine grayish granules. Asci 8-spored, tholus moderately thick, thin when mature.


Ascospores (very) small, (sub-)muriform, cell walls moderately thin to moderately thick, 

endospore thick, in younger ascospores sometimes with thin halo, hyaline, distinctly amyloid, 

sub-globular to oblong with rounded ends, loci predominantly roundish, subglobular to 

oblong with same shaped end cells, transverse septae moderately thick, irregular, 10-30 x 7- 

15(18) µm with 4-6(8) x 1-4(6) loci. Pycnidia present, immersed or in thallus warts, conidia 

bacilliform, c. 2-4 x 0.5 µm. 

Fig. 46. Fibrillithecis halei: growth habit (A, C, D), ascomata (B), section of ascoma margin (E), 

ascospores showing amyloid reaction (F), ascospores (G) and conidia (H). A.: Streimann 61845; 

B., E., G.: FH-lectotype of T. piluliferum; C., F.: FH-lectotype; D.: Weber & McVean s.n.; H.: 

Mangold 36 v. Bar= A: 1.75 mm; B: 0.6 mm; C, D: 1.5 mm; E: 50 µm; F:15 µm; G: 10 µm; H: 6 

µm.


CHEMISTRY – Thallus K-, C-, PD+/- yellow; containing an unknown sterol ['platysporum 

unknown': Rf 57 in B', (dark-)brown after charring with yellowish to greenish aurora in UV] 

(major to minor or lacking), psoromic (major to trace or lacking), 2'-0-demethylpsoromic 

(minor to trace or lacking) acid, and sometimes traces of other unknown substances of the 

psoromic acid chemosyndrome. 

ECOLOGY AND DISTRIBUTION – Fibrillithecis 

halei grows on tree bark and bryophytes in (sub-) 

tropical rainforests and mangroves in altitudes 

ranging from sea level to 1350 m. It is common and 

wide-spread in northern Queensland and New 

South Wales. This pantropical species was reported 

from Hawaii, U.S.A. (Harris, 1990), Venezuela, 

Africa (Frisch, 2006), Réunion (Kalb, 2001), India 

(Nagarkar & al., 1985), Sri Lanka (Hale, 1981), 

Borneo (Hale, 1974, 1981; Sipman, 1993), New 

Caledonia (Hale, 1981). 


corticate thallus, large, usually perithecioid, 

±emergent ascomata, the thick, fibrous proper 

exciple, moderately small, muriform, thick-walled, 

hyaline, amyloid ascospores and the presence of the 

psoromic acid chemosydrome and/or 'platysporum 

unknown'. It is variable in morphology and 

chemistry, while its ascospore and ascomata 


F. halei. 

anatomy is homogenous. In his treatment, which includes the introduction of the new genus 

Fibrillithecis, Frisch (2006) separated three different taxa, the isidiate F. insignis and two 

non-isidiate species: F. vernicosa (containing psoromic acid) and F. platystoma (containing 

'platystoma unknown'). Since chemically intermediate forms, containing both compounds, 

were found in the Australian collections, the latter two names are regarded as conspecific. 

The type collection of Porina halei from Venezuela in FH agrees well with the paleotropic 

material despite ascomata with distinctly free proper exciple and a strongly verrucose thallus 

surface. The latter feature, interpreted as isidia by Harris (1990), can be found in less distinct 

form in several other specimen of F. halei and do not confirm with the 'true' isidia found in F. 

insignis. The detached proper exciple, however, is most likely due to depauperisation of the 

collection. 

SPECIMENS EXAMINED – Australia, Queensland: Lockhart River Settlement road, 36 km SW of Cape 

Weymouth, Streimann 56621 (B, CANB, H). Cooktown, trail from Grassy Hill to Cherry Tree Bay, Lumbsch & 

Guderley 11173i (F). Annan River, Grass Tree Pocket Rd., 38 km S of Cooktown, Streimann 46374 (B, CANB). 

Mt. Windsor, 5 km W of new Forestry Camp, NW of Mossman, Hale 830644, 831876, 831950, 832644 (US). 

Mossman River Gorge, (Hale exsic. as T. diminitum, distrib. Univ. of Colorado Museum, no. 453), Weber & 

McVean s.n. (BM, CANB, COLO, H, MEL, NSW, S, US); Mangold 36 v (F). Fresh Water Gorge, outside of city 

of Cairns, Hale 830746 (US). Fitzroy Island on Great Barrier Reef, 25 km E of Cairns, Aptroot 22280 (ABL). 

Atherton Tablelands: Davies Creek Rd. 17 km S of Kennedy Hwy., S of Davies Creek Falls NP., E of Mareeba, 

Hale 832125 (US); SW of K-1 tree rd. off Palmerston Hwy., 11 km from main hwy. and 2 km N of S.Johnstone 

Forestry Camp, SE of Millaa Millaa, Hale 832413, 832684, 832729 (US); 23 km E of jct. Kennedy Hwy and 

Palmerston Hwy, E of Ravenshoe, Hale 832098 (US). The Boulders, NW of Babinda, S of Cairns, Hale 831371 

(US). 3 km N of Garradunga, Graham Range, N of Innisfail, Hale 831208 (US). State Forest area on Tully Rd., 1 

km from jct. with S. Mission Beach Rd., S of Mission Beach, Hale 831122, 831226, 831754 (US). Dawson 

logging area, State Forest Reserve 605, 24 km S of Koombooloomba turnoff, WSW of Tully, Hale 830607, 

830613 (US). Culpa logging area, 13 km from Koombooloomba rd. turnoff, SE of Tully Falls, Hale 832211 

(US). Edmund Kennedy NP.: On Clift Rd., NW of Cardwell, Hale 832818, 832819 (US); Few km N of


Cardwell, A. & K. Kalb 34285 (hb. Kalb). Kennedy North district, near Meunga Creek, 4 km N of Cardwell, 

Streimann 61845 (B, CANB). Hinchinbrook Isld., northern end, Gilbert 75/797 (HOB). 7.5 km E of Wallaman 

Falls, W of Ingham, Hale 69167, 69171, 832250 (US). Little Crystal Creek Falls, Mt.Spec NP., the falls on 

Paluma Rd., Hale 831204 (US). Cape Hillsborough NP., NW of Mackay, Hale 831705, 832470 (US). New 

South Wales: New England NP., along Tree Fern Gully, Wedin 3591 (UPS). Werrikimbe NP., Beech Plateau, 80 

km NW of Port Macquarie, Streimann 63965 (B, CANB). Burraga Swamp, Allyn River Forest Park, Kantvilas 

s.n. (NSW). Gloucester Tops, Kantvilas s.n. (NSW). Malaysia, Borneo, Sabah, Hale 28344 (US). 

2. 9. 4. Leptotrema Mont. & Bosch, in Miquel, F.A.W., Plantae Junghuhnianae. 4: 483 

(1855). Type species: Leptotrema zollingeri Mont. & Bosch. Type: Java, Zollinger 738 (Lholotype). 

Leptotrema wightii is the only known species in Australia, for a description see there. 

NOTES – This recently resurrected genus (Frisch, 2006) traditionally included taxa with 

brown, muriform ascospores according to Müller’s (1887c) ascospore-based classification. In 

Hale’s revised generic concept (1980, 1981) it was synonymized with Myriotrema based on 

the absence of lateral paraphyses and carbonization. Frisch (2006) only accepts one species, 

L. wightii. In contrast, I accept an additional taxon. The type species L. zollingeri was 

considered conspecific with L. wightii by Frisch (ibid.) but is considered here a distinct 

species based on a different chemistry (see also notes under L. wightii). 

The genus is accepted here tentatively and needs evaluation by molecular data. It is 

distinguished from the similar genus Leucodecton by two characters, the ‘sac-like’ asci 

without a tholus, and ascospores with an early thick-walled stage. Reimnitzia has similar 

ascospores but differs by its Geaster-like ascomata and asci with a ±distinct tholus. The 

differences in exciple morphology (Thelotrema-type proper exciple in Reimnitzia) stated by 

Frisch (ibid.) could not be confirmed, since no lateral paraphyses where found in the 

Reimnitzia specimen examined. Myriotrema is another similar genus, readily distinguished by 

its excipular structure with ±radiating apical hyphae, further also differing in asci with a 

±distinct tholus and ascospores with comparably thin walls in immature stages. Leptotremalike 

asci and ascospores with thick walls from early stages on can also be found in species 

belonging to genera which are otherwise well separated: These include Chapsa lamellifera 

that has ‘Leptotrema-type’ asci and Ocellularia bahiana with similar ascospores. Other 

specific characters for Leptotrema include a basal thallus layer of conglutinated hyphae, 

which also occurs in R. santensis, and the columnar arrangement of calcium oxalate crystals 

that occurs in several other species (e.g. R. santensis, Myriotrema phaeosporum, Thelotrema 

myriocarpum, Leucodecton compunctellum). 


Leptotrema wightii (Tayl.) Müll. Arg. 

Flora 65: 499 (1882). Bas.: Endocarpon wightii Tayl., Lond. J. Bot. 6: 155 (1847). Thelotrema wightii 

(Tayl.) Nyl., Mém. Soc. Sci. Cherbourg 5: 118 (1857). Phaeotrema wightii (Tayl.) Zahlbr., Ark. Bot. 31A(1): 48 

(1944). Myriotrema wightii (Tayl.) Hale, Mycotaxon 11: 135 (1980). Type: India, Madras, Wight s.n. (FH- 

Tayl.!-lectotype, selected by Hale [1974b: 43]; BM!-, G-isolectotypes). 

Endocarpon baileyi Stirt., Trans. Proc. Roy. Soc. Vict. 17: 74 (1881). - Leptotrema baileyi (Stirt.) Shirl., 

Proc. Roy. Soc. Queensl. 6: 194 (1889). Type: Australia, Queensland, Brisbane, 1878, Bailey 249 (GLAMholotype; 

BRI!-isotype).


Thelotrema ravenelii Tuck., Amer. J. Arts and Sci, ser. 2, 25: 426 (1858). Leptotrema ravenelii (Tuck.) Fink, 

Lichen Flora U.S.: 133 (1935). Type: U.S.A., South Carolina, Ravenel 151 (FH-Tuck.-lectotype, selected by 

Salisbury [1971b: 35]; NY!-, US!-isolectotypes). 

Thelotrema subconcretum Leight., Trans. Linn. Soc. London 27: 169 (1869). Phaeotrema subconcretum 

(Leight.) Müll. Arg.: Mém. Soc. Phys. Hist. Nat. Genève 29(8): 10 (1887c). Leptotrema subconcretum (Leight.) 

Müll. Arg., Nuov. Giorn. Bot. Ital. 23: 277 (1891a). Type: Sri Lanka, Central Province, Thwaites 89 (BM!lectotype, 

selected by Hale [1974b: 43]). 

[For additional synonymy see Frisch (2006), see also Mangold & al. (2006) regarding the synonymy of 

Thelotrema leiospodium Nyl. and the note below regarding L. zollingeri Mont. & Bosch.] 


Fig. 48. Leptotrema wightii: growth habit (A), thallus section showing anthraquinone crystal cluster 

(arrow) (B), ascomata (C), ascoma and thallus section (D) and ascospores (E). A.: BRI–isotype of 

E. baileyi; B., E.: Elix 35434; C., D.: Hale 831324. Bar= A: 8 mm; B: 1.3 mm; C: 1 mm; D: 200 

µm; E: 14 µm. 

Thallus epiphloedal, thick to very thick, usually bulging and flaking away from the 

substrate, up to c. 1000 µm high, pale greenish gray. Surface dull, smooth, continuous to 

rarely rimose, with a bright reticulate structure, forming a grainy-speckled pattern, unfissured, 

sometimes coarsely cracked. True cortex absent, thallus covered by a ±continuous protocortex 

up to c. 20 µm thick. Algal layer well developed and continuous, becoming incontinuous due 

to crystal inclusions, calcium oxalate crystals abundant, large, clustered, forming columnar 

structures, medulla with conspicuous, frequent or rarely infrequent bright red anthraquinone 

crystals. Basal thallus sometimes formed of strongly conglutinated hyphae, forming a lower 

cortex-like structure. Vegetative propagules not seen. Ascomata inconspicuous, small to 

moderately large, up to c. 600 µm in diam., ±roundish, peri- to apothecioid, predominantly 

solitary, immersed. Disc usually not visible from surface, rarely becoming partly visible, pale 

to dark brown, epruinose. Pores small to wide to rarely gaping, up to c. 400 µm in diam., in 

mature ascomata formed by proper exciple, roundish to rarely slightly irregular, with entire 

margin, apical proper exciple becoming ±visible from the outside, forming a fused to 

somewhat (rarely distinctly) free inner pore margin, incurved and usually ±sunken, bright-


translucent to off-white. Thalline rim margin thick, entire, with same form as pore margin, 

often brighter than thallus. Proper exciple predominantly fused to slightly detached, only in 

old, gaping ascomata becoming distinctly free, (moderately) thick, hyaline internally to 

yellowish or orange-brown marginally, sometimes with grayish granules incorporated, nonamyloid. 

Hymenium up to c. 250 µm high, non-inspersed, strongly conglutinated, paraphyses 

straight to slightly bent, parallel to somewhat interwoven, slightly branched towards the 

apical hymenium, with distinctly thickened tips, lateral paraphyses and columellar structures 

absent. Epihymenium moderately thin, hyaline, sometimes with fine grayish granules. Asci 8spored, 

tholus or distinctly thickened parts absent. Ascospores (very) small, (sub-)muriform, 

cell walls thick in younger stages, in mature ascospores cell walls moderately thick to 

moderately thin, endospore (moderately) thin, non-halonate, becoming brown at early 

maturity, non-amyloid to faintly amyloid, subglobose to oblong to more rarely ellipsoid, with 

predominantly rounded ends, loci roundish to (slightly) angular, predominantly irregular, 

septae becoming (moderately) thin, irregular or often with one central, more distinct septum, 

10-30 x 8-15 µm, with 3-6 x 1-4 loci. Pycnidia not seen, fide Frisch (2006) immersed or in 

thallus warts, conidia bacilliform, 5-7 x 1-1.2 µm. 

CHEMISTRY – Thallus K- with K+ purple crystals, C-, PD-; containing an unknown 

anthraquinone (RF-values 33/14/14 in solvent systems A/B'/C). 

ECOLOGY AND DISTRIBUTION – Leptotrema 

wightii was collected in Australia on bark of trees 

or shrubs, often overgrowing adjacent mosses. It 

was most frequently found in seasonally wet areas 

such as dry monsoon shrubs and forests mainly at 

river flats; rarely in rainforests. It occurs in tropical 

to warm-temperate climates in altitudes ranging 

from sea level to 1000 m. It is a common and widespread 

species, occurring throughout Queensland, 

northern and southern-central New South Wales. In 

Queensland it often extends somewhat into the 

hinterland. Besides Australia it was reported from 

Hawaii (Magnusson & Zahlbruckner, 1944), the 

Neo-tropics (Harris, 1990; Hale, 1974; Redinger, 

1936), Africa (Frisch, 2006), India, Sri Lanka, 

Philippines (Hale, 1981), Sarawak (as T. forminulosum, 

Krempelhuber, 1875), indicating a pan(sub)tropical 

distribution that extends into warmtemperate 

climates. 


L. wightii. 

NOTES – This taxon can be easily identified by the thick, bulging thallus with reticulate 

surface and conspicuous red anthraquinone crystals, immersed ascomata and small, (sub-) 

muriform, roundish, brown, non-amyloid ascospores with an irregular loci arrangement. A 

very similar species in Australia is M. phaeosporum, for differences see under that species. 

Thelotrema leiospodium, known from Europe and Macaronesia, was long treated as a 

synonym (Salisbury, 1971) but can be readily distinguished by the absence of anthraquinone 

crystals, the distinct lateral paraphyses and larger (up to 40 µm long, with up to 10 x 5 loci), 

fusiform ascospores. [See Mangold & al. (2006) for a more detailed discussion, see also there 

(ibid.) for a discussion of the taxonomic treatment of the similar Leptotrema lithophila Oxner 

described from far-east Asia.] The examined isotype of L. zollingeri in US (Java, Zollinger 

738 - holotype in L) contains the hypoprotocetraric acid chemosyndrome and lacks


anthraquinone crystals. This taxon, listed as a synonym of L. wightii by Frisch (2006), is thus 

here regarded as a distinct species. 

SPECIMENS EXAMINED – Uncertain location: Woolston (?New Zealand), Wilson s.n. (NSW-539379). 

Australia, Queensland: Bloomfield River, 55 km SSE of Cooktown, Streimann 57282 (B, CANB). Royal Arch 

Caves NP., 5 km SW of Chillagoe, Streimann 46481 (CANB). Forty Mile Shrub NP., 53 km E of Mount 

Surprise, Streimann 46720 (CANB). Conway Range NP., near Shute Harbour-Airlie Beach, Hale 831935 (US). 

Bunya Mountains NP.: 2 km E of Munro's Camp, Scott 705 (BRI); Track from Dandabah to Pine George 

Lookout via Festoon Falls, Lumbsch 10998i (F); Just before leaving NP. on the N, Hale 831324 (US). Port 

Curtis, Rockhampton, Thozet 755 (G, M, MEL). Goodnight Shrub near Gympie, Ryan s.n. (BRI-689506). 

Moreton, Rosewood Shrub, Bailey s.n. (BRI-689507, MEL-5792). Booloumba Creek SF., SW of Kenilworth, 

Hale 831285 (US). Maroochy, Bailey 811 (BRI). 14 km WSW of Monto, Streimann 9864 (CANB). Kalpowar 

Forest Drive, c. 40 km NE of Monto, Hale 831183 (US). Mt. Mee SF., 6 km NW of Forestry Office, Hale 

830853 (US). Hurdle Gully, Coominglah SF., Elix 35434 (CANB). Isla Gorge NP., 27 km NNE of Taroom, Elix 

35133 (CANB). Pine Mountain SF., near Flutter Creek, Elix 34790 (B, CANB). Between Springsure and 

Emerald, McVean 6396 (COLO). Mt. Glorious near Brisbane, Rogers s.n. (BRI-689027). Mt. Beppo, near Esk, 

Rogers s.n. (BRI-689502). Lowood, Wilson s.n. (NSW-539381). 10.5 km SE of Toogoolawah, 2 km SE of Mt. 

Beppo, Tibell 12530 (UPS). Brisbane, Bailey s.n. (G-10194/29-31). Lingalonga, Flagstone Creek near 

Toowoomba, Swarbrick 694630 (BRI). Darling Downs, Toowoomba, Hartmann s.n. (G, MEL-26203). 

Cunninghams Gap NP., c. 40 km NW of Warwick, A. & K. Kalb 21409 (hb. Kalb). Killarney, Wilson s.n. 

(NSW-539383). New South Wales: Wollongbar, Wilson s.n. (NSW-539382.A). Hillend, Wilson s.n. (NSW- 

539385). Uncertain location, ‘Shirley book’, p. 23, s.c. (BRI-AQ721246); ‘Shirley book’, p. 'D', s.c. (BRI- 

AQ721713). 

2. 9. 5. Leucodecton Massal., Atti I. Reale Istit. Veneto, ser. 3, 5: 325 (1860). Type species: 

Leucodecton compunctum (Ach.) Massal. Type: India Occidentalis, on Cinchona 

angustifolia, s.c. (H-Ach.734!-lectotype, selected by Frisch [2006: 148]; UPSisolectotype). 

Enterostigma Müll. Arg., Flora 68: 254 (1885) [nom. superfl. pro Leucodecton Massal.]. Chiodecton subgen. 

Enterostigma (Müll. Arg.) Vain., Ètude Lich. Brésil 2: 138 (1890). Enterostigmatomyces Cif. & Thomas., Atti 

Ist. Bot. Lab. Crittog. Univ. Pavia, ser. 5, vol. 10: 73 (1953). Type species: Enterostigma compunctum (Ach.) 

Müll. Arg. [=Leucodecton compunctum (Ach.) Massal]. 

THALLUS – Crustose, corticolous, epi- to hypophloedal, moderately thin to moderately 

thick, up to c. 400-500 µm high, rarely up to 1 mm high. Mostly pale, in shades of gray or 

green with olive, yellowish or whitish tones, sometimes with grainy-speckled pattern. Surface 

dull to slightly shiny, smooth to ±farinose or rough, continuous to rugose or verrucose to 

verruculose, unfissured to fissured or more rarely rimose to areolate. Prothallus thin to 

indistinct, brown. Predominantly ecorticate, sometimes without cortical structures to usually 

without distinct true cortex and covered by a up to c. 10-30 µm thick, continuous to 

discontinuous protocortex, very rarely with distinctly conglutinated parts forming a partial 

true cortex consisting of irregular to rarely periclinal hyphae. Algal layer continuous and well 

developed to more rarely discontinuous and poorly developed, calcium oxalate crystals 

predominantly abundant, small to large, scattered to clustered, sometimes forming columnar 

structures, rarely forming layers in lower thallus (L. subcompunctum). Distinct medulla layer 

mostly absent, rarely present (L. glaucescens). Vegetative propagules not seen, soralia 

occurring in L. sorediiferum (not known from Australia). 

ASCOMATA – Predominantly inconspicuous to rarely conspicuous, small to moderately 

large, c. up to 400-600 µm in diam., usually roundish to slightly irregular, sometimes 

distinctly irregular or angular . Predominantly apothecioid to rarely perithecioid, solitary to 

marginally fused, sometimes also distinctly fused and clustered, forming stroma-like 

structures, non-regenerating, predominantly immersed to more rarely slightly raised to very 

rarely distinctly emergent, then hemispherical with same surface as thallus. Disc rarely not


visible to more often partly or sometimes entirely visible from surface, grayish, distinctly 

pruinose. Pores small or moderately wide, rarely wide to gaping, c. up to 50-550 µm in diam., 

formed by proper exciple, roundish to irregular or angular, pore margin predominantly entire 

to more rarely slightly split or dentate. Proper exciple apically or in upper parts becoming 

visible from surface, usually distinctly free to rarely somewhat fused, particularly in younger 

stages, often conspicuously short, mostly whitish to off-white, more rarely grayish or 

brownish, sometimes shrunken, incurved to erect to more rarely somewhat recurved. Thalline 

rim margin moderately thin to moderately thick, roundish to slightly irregular, more rarely 

distinctly irregular or angular, predominantly wide to gaping, rarely small, entire to rarely 

slightly eroded, concolorous to ±brighter than thallus, in emergent ascomata incurved. Proper 

exciple usually apically or in upper parts free, at least in older stages, rarely thin to 

moderately thick, hyaline to pale yellowish or rarely pale brownish internally, yellowish or 

brownish, more rarely with orange or grayish tones marginally, apically sometimes darkishbrown 

to slightly carbonized, non-amyloid to more rarely faintly amyloid at the base. 

Subhymenium indistinct, thin and with same color as basal exciple. Hymenium non-amyloid, 

moderately to distinctly cupular, up to c. 120-200 µm high, non-inspersed, clear, moderately 

to strongly conglutinated. Paraphyses straight to ±bent, sometimes distinctly curly in apical 

parts, distinctly interwoven, predominantly slightly branched towards the margins, tips 

slightly to rarely distinctly thickened, rarely somewhat irregular. Lateral paraphyses and true 

columella absent, in fused ascomata sometimes columella-like structures present. 

Epihymenium hyaline, predominantly moderately thin to thick with grayish to brownish 

granules and often small crystals. 

Asci predominantly 8-spored, rarely 1-4-spored, non-amyloid, clavate, ascus walls mostly 

not thickened and with moderately thin to thick, in mature asci thin to invisible tholus, 

sometimes younger asci without distinct tholus and with thickened ascus walls throughout, at 

maturity evenly thinning or thinning in lower parts only (L. compunctellum and L. occultum). 

Ascospores uni- to biseriate, very small to large, 7-130 x 5-35 µm, predominantly 

submuriform, more rarely eumuriform or transversely septate. Cell walls mostly moderately 

thick to thick, predominantly smooth to rarely crenate or irregular at late maturity, endospore 

in muriform ascospores thin to moderately thick, non-halonate, predominantly brown and 

non-amyloid to faintly amyloid, in L. compunctum (not known from Australia) hyaline to 

yellowish in depauperate ascospores and strongly amyloid (see also notes). Predominantly 

oblong to ellipsoid, more rarely subglobose or fusiform, transversely septate ascospores fusito 

claviform, with roundish to narrowed-roundish to more rarely subacute, very rarely 

distinctly acute ends; with 4-45 x 0-12 loci, loci roundish to angular, subglobose to lentiform 

or ±irregular, with same shaped to rarely hemispherical or conical end cells, transverse septae 

thin to moderately thick. 

PYCNIDIA – Found in several species immersed with dark pore area, conidia variable, 

obovate to fusi- or bacilliform, up to c. 3-6 x 1 µm. 

CHEMISTRY – Secondary compounds present to rarely absent (L. compunctum), 

predominantly of the stictic acid, rarely norstictic acid chemosyndrome. 

ECOLOGY AND DISTRIBUTION – In Australia, Leucodecton species grow on tree bark 

predominantly in (sub)tropical to rarely warm-temperate rainforests and wet sclerophyll 

forests, more rarely in tropical mangroves, in altitudes ranging between sea level and 1500 m. 

They occur in north-western Northern Territory, and in Queensland to central New South 

Wales and Lord Howe Island. Four of the five currently known Leucodecton species 

occurring in Australia are pantropical, ±extending into subtropical or warm-temperate zones; 

L. albidulum is thus far only known in Australasia.


N OTES –This neglected genus was described by Massalongo (1860) and recently 

resurrected (Frisch, 2006). 

The species in Leucodecton are mainly characterized by a predominantly ecorticate thallus, 

apothecioid ascomata with ±free proper exciple that consists of distinctly 

paraplectenchymatous hyphae without a parallel (basally) or radiating (apically) orientation 

and no distinct carbonization (“Leucodecton-type”), distinctly interwoven, often slightly 

branched paraphyses, and absence of lateral paraphyses. Further, most of the Leucodecton 

taxa have ±distinctly muriform, brown, non- to faintly amyloid ascospores and contain stictic 

or norstictic acid. Exceptions are L. albidulum with transversely septate ascospores and L. 

compunctum with hyaline, distinctly amyloid ascospores and no secondary metabolites. 

The most similar genus is Leptotrema, for differences see under that genus. Also similar 

are members of Myriotrema, this genus is mainly distinguished by a different exciple type 

with parallel hyphae and radiating tips (‘Ocellularia-type’). In particular Myriotrema species 

with brown ascospores and the stictic acid chemosyndrome are difficult to distinguish, see at 

Myriotrema. 

Leucodecton forms a well supported clade in the molecular analysis of Frisch & al. (2006) 

and its distinction from Myriotrema (see above) is supported. 


Leucodecton albidulum (Nyl.) Mangold comb. nov. ined. 

Bas.: Thelotrema albidulum Nyl., Annal. Scienc. Nat. Bot. 4(15): 46 (1861). Myriotrema albidulum (Nyl.) 

Hale, Mycotaxon 11: 132 (1980). Type: Nova Caledonia, Balade, 1867, Vieillard s.n. (H-Nyl. 22767!-holotype; 

H-Nyl. 22770!-, PC-isotypes). 


Thallus hypo- to predominantly epiphloedal, moderately thick, up to c. 500 µm high, offwhite 

to (pale) grayish-green. Surface dull, smooth to more often finely pruinose, continuous 

to sometimes verrucose, ±distinctly fissured. Cortex structures absent. Algal layer well to 

sometimes poorly developed, continuous to incontinuous, calcium oxalate crystals abundant 

to more rarely sparse, scattered or clustered. Vegetative propagules not seen. Ascomata 

inconspicuous, moderately small to sometimes moderately large, up to c. 600 µm in diam., 

roundish to more often ±irregular, apothecioid, solitary to fused and clustered, sometimes 

forming stroma-like structures, predominantly immersed. Disc predominantly entirely visible 

from surface, grayish, pruinose. Pores wide to gaping, up to c. 550 µm in diam., formed by 

proper exciple, proper exciple apically visible from surface, free, rarely roundish to more 

often ±irregular or angular, predominantly entire, very short, whitish to off-white, often 

±shrunken, erect to slightly incurved. Thalline rim margin moderately thin to moderately 

thick, roundish to roundish-irregular or angular, gaping, entire to slightly eroded, brighter 

than thallus or concolorous with thallus. Exciple becoming entirely free, thin, pale yellowish 

internally to yellowish-brown or brownish marginally, non-amyloid. Hymenium up to c. 120 

µm high, non-inspersed, moderately conglutinated, paraphyses straight to slightly bent, 

distinctly interwoven, slightly branched towards the margins, lateral paraphyses and true 

columella absent, in fused ascomata sometimes with columella-like structures. Epihymenium 

(moderately) thick, with grayish-brown granules. Asci 8-spored, tholus (moderately) thick, 

thin when mature. Ascospores (moderately) small, transversely septate, cell walls moderately 

thin to moderately thick, becoming ±distinctly crenate to irregular at late maturity, endospore


becoming (moderately) thick, non-halonate, brown, ±faintly amyloid, predominantly fusi- to 

claviform, sometimes slightly bent, with roundish to more often (sub)acute ends, loci angular 

in immature ascospores, becoming roundish and subglobose to slightly lentiform with 

maturity, end cells hemispherical to more often conical, septae moderately thin to moderately 

thick, regular, 25-45 x 5-7 µm with 8-12 loci. Pycnidia present, immersed with darkened pore 

area, conidia bacilliform, up to c. 6 x 1 µm. 


constictic, hypostictic (major to minor), α-acetyl-constictic, hypoconstictic and cryptostictic 

(traces) acids. 

Fig. 50. Leucodecton albidulum: growth habit (A), ascomata (B), mature ascospores (C), immature 

ascospore (D), pycnidia (E) and conidia (F). A.: Wilson s.n. (NSW); B., D.: Lumbsch & Mangold 

19116 p; C.: H-lectotype; E., F.: Mangold 35 e. Bar= A: 2 mm; B: 1 mm; C: 7.5 µm; D: 5 µm. E: 

200 µm, F: 6 µm.


ECOLOGY AND DISTRIBUTION – Leucodecton 

albidulum was collected in Australia on tree bark in 

(sub)tropical rainforests in altitudes ranging from 

200 to 900 m. It is a rare but widely distributed 

species occurring in northern and northern-central 

Queensland and in the Queensland/New South 

Wales border region. This is the first report for 

Australia, otherwise it is only known from the type 

location New Caledonia. 

NOTES – This taxon is readily characterized by 

the thick, ecorticate thallus, the ±distinctly angular, 

clustered ascomata with stroma-like arrangement, 

the branched and interwoven hymenium the 

transversely septate, brown, narrow ascospores 

with (sub-)acute ends and the presence of the stictic 

acid chemosydrome. Leucodecton samaranum 

(Vain.) Mangold comb. nov. ined. 7 from the 

Philippines is morphologically almost identical 

with L. albidulum, and agrees in containing the 

stictic acid chemosydrome but has distinctly smaller ascospores (15-20 x 4-6 µm with 4-5 

loci). A similar species from Australia is L. glaucescens, which can be readily distinguished 

by smaller, submuriform ascospores. This is the first known member of Leucodecton with 

transversely septate ascospores and bacilliform conidia. However, regarding the thallus and 

ascoma anatomy, as well as the chemistry, the taxon is in full accordance with other members 

in this genus. 

SPECIMENS EXAMINED – Australia, Queensland: Daintree NP., Mossman Gorge Section, near Rex Creek 

Swing Bridge, Mangold 35 e (F). Atherton Tablelands: Davies Creek Rd. 17 km S of Kennedy Hwy., E of 

Mareeba, Hale 832335 (US); Lake Euranoo, Hale 831742 (US), Mangold 38 e (F); Danbullah Forest Drive, 1 

km W of Cathedral Fig Tree, Hale 831570 (US); Along west boundary of Lake Eacham NP., Hale 831229 (US); 

SW of K-1 tree rd. off Palmerston Hwy., 2 km N of S.Johnstone Forestry Camp, SE of Millaa Millaa, Hale 

831228 (US). Eungella NP.: "Palm Walk", 2.5 km SE of Eungella, A. & K. Kalb 34249, 34264 (hb. Kalb); Near 

Pease's Lookout, Hale 831548 (US); Finch Hatton Gorge, Lumbsch & Mangold 19116 p (F). Mary Cairncross 

Park, SE of Maleny, Hale 831224, 831578, 831728 (US). New South Wales: Mount Warning NP., track from 

parking lot to Lyrebird Lookout, Mangold 17 g (F). Big Shrub Flora Reserve, Night Cap Forest Drive, W of 

Mullumbimby, Hale 831396 (US). Big Shrub, Richmond River, Jun. 1894, Wilson s.n. (MEL, NSW, US). 

Leucodecton compunctellum (Nyl.) A. Frisch 


L. albidulum. 

Bibl. Lichenol. 92: 155 (2006). Bas.: Thelotrema compunctellum Nyl., Bull. Soc. Linn. Normand ser. 2, 2: 77 

(1868). Leptotrema compunctellum (Nyl.) Zahlbr., Cat. Lich. Univ. II: 632 (1923). Type: New Caledonia, 

Loyalty, 1864, Thiébault s.n. (H-Nyl. 22703!-lectotype, selected by Hale [1972 in herb.]). 

Thelotrema monosporum var. subgemium Nyl., Bull. Soc. Linn. Normand. 2, 2: 77 (1868). Thelotrema 

subgemium Nyl., Sertum Lichen. Tropic. Labuan et Signapore, p. 5 (1891). Type: New Caledonia, Lifu 

(Loyalty), 1864, Thiebaut s.n. (H-Nyl. 22702!-lectotype, selected by Hale [1972 in herb.]). 

Thelotrema elachistoteron Leight., Trans. Linn. Soc. Lond. 27: 169 (1870). Leptotrema elachistoteron 

(Leight.) Patw. & Kulk., Norw. J. Bot., 24: 128 (1977). Myriotrema elachistoteron (Leight.) Hale, Mycotaxon 

11: 133 (1980). Leucodecton elachistoteron (Leight.) Frisch, Bibl. Lichen. 92: 155 (2006). Type: Sri Lanka, 

Central Province, Thwaites C.L. 132 (BM!–lectotype, selected by Hale [1981: 278]). 

7 Leucodecton samaranum (Vain.) Mangold comb. nov. ined., Bas.: Thelotrema samaranum Vain., Annal. 

Acad. Scient. Fennic. ser. A 15(6): 185 (1921). Phaeotrema samaranum (Vain.) Zahlbr., Cat. Lich. Univ. II: 610 

(1923). Type: Philippines, Samar, Catubig River, Bur. Sci. 24931 pr.p., Feb.-Mar. 1916, Edano s.n. (TUR-Vain. 

31266!-holotype).


Thelotrema reclusum Kremp. in Nyl., Bull. Soc. Linn. Normand., 7(2): 168 (1873). Leptotrema reclusum 

(Kremp.) Zahlbr., Cat. Lich. Univ. II: 639 (1923). Myriotrema reclusum (Kremp.) Hale, Mycotaxon 11: 135 

(1980). Type: Andaman Islands, Kurz 21 (M–lectotype, selected by Hale [1978: 54]; BM!-, FH-Tuck.-, H-Nyl.-, 

UPS-, W-, ZT–isolectotypes). 

Anthracothecium oligosporum Müll.Arg., Flora 71: 48 (1888). Leptotrema oligosporum (Müll. Arg.) Patw. & 

Makhija, Bryologist 83: 368 (1980). Type: Australia, Queensland, Lower Herbert River (near Ingham), Wickham 

s.n. (G!–lectotype, selected by Patwardhan & Makhija [1980: 368]). 

Thelotrema microglaenoides Vain., J. Bot. 34: 206 (1896) [non Thelotrema microglaenoides Vain., Ann. 

Univ. fenn. åbo, ser. A, 2(3): 32 (1926) = nom. illegit.! {=Diploschistes diploschistoides (Vain.) Salisb. fide 

Guderley & Lumbsch (1996)}]. Leptotrema microglaenoides (Vain.) Zahlbr., Cat. Lich. Univ. II: 637 (1923). 

Type: St. Vincent, Elliott 266 (TUR-Vain.!–holotype, BM–isotype). 

Leptotrema deceptum Hale, Sm. Contr. Bot. 16: 39 (1974). Myriotrema deceptum (Hale) Hale, Mycotaxon 

11: 133 (1980). Type: Dominica, Dec.1971, Hale 37860 (US!–holotype). 

Myriotrema nuwarense Hale, Bull. Br. Mus. Nat. Hist. (Bot.) 8(3): 289 (1981). Leptotrema nuwarense (Hale) 

Nagark., Sethy & Patwardhan, Mycotaxon 27: 74 (1986). Leucodecton nuwarense (Hale) Frisch, Bibl. Lichen. 

92: 155 (2006). Type: Sri Lanka, Central Province, Nuwara Eliya District, Hale 50272 (US!–holotype, 

BM–isotype). 

Leucodecton biokense A. Frisch, Bibl. Lichen. 92: 152 (2006). Type: Equatorial Guinea, Bioko, Mann s.n. 

(BM-holotype, BM!-isotype). 


Thallus epi- to hypophloedal, variable in thickness, thin to moderately thick, up to c. 500 

µm high, pale gray to greenish gray or pale yellowish-brown, often with grainy-speckled 

pattern. Surface dull to slightly shiny, smooth, continuous to rugose and/or minutely 

verruculose, unfissured to fissured. True cortex absent, thallus predominantly covered by an 

±continuous protocortex up to c. 20 µm thick. Algal layer poorly to well developed, 

predominantly continuous, calcium oxalate crystals usually abundant, large and clustered, 

sometimes forming columnar structures. Vegetative propagules not seen. Ascomata 

predominantly inconspicuous, small to moderately large, up to c. 400 µm in diam., roundish, 

peri- to rarely somewhat apothecioid, solitary, immersed to slightly emergent, then 

hemispherical with same surface as thallus. Disc not visible from surface. Pores tiny to more 

rarely small, up to c. 50(100) µm in diam., roundish with entire margin, apical proper exciple 

becoming visible from the outside, forming a fused to ±free inner pore margin, incurved, 

bright-translucent to pale brownish, in older ascomata often thinning, then appearing more 

dark. Thalline rim margin moderately thick, roundish, entire, (moderately) small, incurved, 

concolorous to sometimes brighter than thallus. Proper exciple fused to apically free in older 

ascomata, moderately thin to moderately thick, hyaline to pale yellowish internally, 

(pale)orange- to yellowish-brown or brownish marginally, non-amyloid or slightly amyloid at 

the base. Hymenium up to c. 200 µm high, non-inspersed, strongly conglutinated, paraphyses 

±bent to often distinctly curly in apical parts, distinctly interwoven, unbranched to slightly 

branched towards the margins, with slightly thickened tips, lateral paraphyses and columellar 

structures absent. Epihymenium indistinct or lacking, hyaline and without granules in 

perithecioid ascomata, in apothecioid ascomata thin and with grayish-brown granules and 

small crystals. Asci 1-4-spored, distinct tholus usually not developed or thin, in younger asci 

often with moderately thickened walls throughout, thinning in all parts at maturity. 

Ascospores (moderately) large, eumuriform, cell walls and endospore moderately thick, nonhalonate, 

becoming distinctly brown at late stage of maturity, non-amyloid to faintly amyloid, 

predominantly only in younger ascospores, oblong to oblong-ellipsoid with rounded to 

narrowed-rounded ends, loci roundish to angular, predominantly irregular, transverse septae 

thin but distinct, regular, 35-80 x 10-35 µm in 2- to 4-spored asci, 60-130 x 10-35 µm in 

single-spored asci, with 10-45 x 3-12 loci. Pycnidia not seen.



(majors), cryptostictic, hypoconstictic and hypostictic (minors to traces) acids. 

Fig. 52. Leucodecton compunctelum: growth habit (A), ascomata (B, C), ascoma and thallus section 

(D) mature ascospores (E, G, H) and immature ascospore (F). A.: US–holotype of M. nuwarense; 

B.: Hale 831086; C., E.: Hale 830870; D., H.: BM-lectotype of T. elachistoteron; F.: US-holotype 

of L. deceptum; G.: H-lectotype of T. monosporum var. subgemium. Bar= A: 2 mm; B, C: 1 mm; 

D: 100 µm; E, F: 15 µm; G: 30 µm; H: 20 µm. 

ECOLOGY AND DISTRIBUTION – Leucodecton compunctellum occurs on tree bark, predominantly 

in moist lowland habitats such as tropical mangroves and (sub)tropical coastal 

rainforests, more rarely in (sub)tropical montane rainforests, in altitudes ranging from sea 

level to 1100 m. It is common and wide-spread in the north-western Northern Territory and 

Queensland. The pantropical species is also known from the Neotropics (Hale, 1974, 1978; 

Harris, 1995), Africa, India (Patw. & Kulkarni, 1977), Sri Lanka, Andaman Islands and New 

Caledonia.


NOTES – This taxon is characterized by the the 

(moderately) large, eumuriform, brown, thickwalled 

ascospores and the presence of the stictic 

acid chemosyndrome. It is variable, however, 

intermediate stages of all characters, such as 

thickness and coloration of the thallus, size of 

ascospores and number per ascus were found (see 

also Mangold & al., 2006). Several similar, stictic 

acid containing species are known from Australia, 

which all differ in smaller ascospores including L. 

subcompunctum, M. desquamans, M. phaeosporum 

and M. trypaneoides. Leucodecton bisporum (Nyl.) 

Mangold comb. nov. ined. 8 from Guadeloupe is 

another similar species, however, the observed type 

material in H-Nyl. differs in having a distinctly 

inspersed hymenium and larger ascospores (up to 

180 µm long). 

SPECIMENS EXAMINED – Australia, Northern Territory: 

Palmerston, 20 km SE of Darwin, A. & K. Kalb 30748, 30754 

(hb. Kalb). Queensland: Cape York Peninsula, Tozers Gap, Iron Range NP., Streimann 56352 (CANB). Cape 

Tribulation Area: C. 15 km E of Daintree, Lumbsch & Mangold 19166 n, 19167 f (F); Between Pilgrim Sands 

and Thornton Beach, Lumbsch & Mangold 19160 e, j, r, t, z, 19161 a, c, e, g, y, hb, va, 19162 l, ob (F); Myall 

Beach, Mangold 31 i, n (F); Walking track from Myall Beach to Cape Tribulation Beach, Mangold 32 e, l, q (F); 

N of Daintree, Hale 831086 (US); Cape Tribulation Beach, Hale 832776 (US). Mt. Windsor, NW of Mossman, 

Hale 830870, 830874, 830989, 832113, 832226 (US). Daintree NP., Mossman Gorge Sct., 26 km NNE of 

Mossman, Streimann 46018 (CANB). Upper Mossman River, Mangold 35 x (F). Cooga Beach, c. 5 km E of 

Mossman, Lumbsch & Mangold 19169 g, h, i (F). W of Palm Cove, c. 25 km N of Cairns, A. & K. Kalb 19942 

(hb. Kalb). Machans Beach, N of Cairns, A. & K. Kalb 21155, 21161, 21185 (hb. Kalb). Atherton Tablelands: 

Kuranda Range, SE of Kuranda, A. & K. Kalb 19902 (hb. Kalb); Stallion Pocket logging area, S of Gordonvale, 

Hale 832435 (US); 15 km along Mulgrave River Rd. from intersection with the Gordonvale Rd., SW of 

Gordonvale, Hale 830726, 831088 (US); Mt. Chalmynia logging area, 15 km from Bruce Hwy, Hale 830993, 

832209 (US). Babinda Boulders, Mangold 39 o (F). Francis Range, Woopen Creek Rd., Hale 832770 (US). 

Edmund Kennedy NP., NW of Cardwell, Hale 832170, 832717 (US). Conway Peninsula, E of Proserpine, Scott 

618 (BRI). Conway Range NP., near Shute Harbour-Airlie Beach, Hale 831258, 832112, 832264 (US). Cape 

Hillsborough NP., NW of Mackay, Hale 830278, 831468 (US). Fraser Coast, River Heads, Lumbsch & Mangold 

19092 q (F). Blackall Ranges (Nambour area), Wilson 2041 pr.p. (NSW). Guadeloupe, Chemin des Mamelles, 

Hale 31382 [as T. bisporum] (US). Dominica, Hale 38139 (US). Sri Lanka, Hale 50170 (US). 

Leucodecton glaucescens (Nyl.) A. Frisch 

Bibl. Lichenol. 92: 164 (2006). Bas.: Thelotrema glaucescens Nyl., Ann. Sci. Nat. Bot. 4(19): 332 (1863b). 

Leptotrema glaucescens (Nyl.) Müll.Arg., Flora 65: 499 (1882). Myriotrema glaucescens (Nyl.) Hale, 

Mycotaxon 11: 133 (1980). Type: U.S.A., Louisiana, Dr. Hale s.n. (FH-Tuck.-lectotype, selected by Hale [1981: 

282]; H-Nyl.-isolectotype). 



L. compunctellum. 

8 Leucodecton bisporum (Nyl.) Mangold comb. nov. ined. Bas.: Thelotrema bisporum Nyl. apud Hue, Nouv. 

Archiv. du Museum, 3(3): 96 (1891) [ex nomen nudum: Thelotrema bisporum Nyl., Mem. Soc. Sci. Nat. 

Cherbourg 5: 118 (1857)]. Type: Guadeloupe, L'Herminier s.n. (H-Nyl. 22489!-lectotype, selected by Hale 

[1972, in herb.]).


Fig. 54. Leucodecton glaucescens: growth habit (A-C), ascomata (D) and ascospores (E, F). A., D., 

E., F.: Lumbsch & Mangold 19117 g; B.: Hale 58568; C.: Hale 832559. Bar= A: 2 mm; B: 1.5 

mm; C: 2.5 mm; D: 500 µm; E: 6 µm; F: 3.5 µm. 

Thallus epiphloedal, thick to very thick, up to c. 1 mm high, bulging and partly flaking 

away from substratum, grayish to grayish-green or pale yellowish. Surface dull to rarely 

slightly shiny, smooth to somewhat mealy, often with speckled pattern, continuous to 

verruculose, unfissured to fissured or rimose. True cortex absent, thallus covered by 

±continuous protocortex up to c. 25 µm thick. Algal layer usually continuous and well 

developed, sometimes becoming incontinuous due to crystal inclusions, calcium oxalate 

crystals abundant, small to large, scattered or clustered, sometimes forming column-like 

structures, distinct, well developed medulla layer present. Vegetative propagules not seen.


Ascomata abundant, conspicuous, moderately small to moderately large, up to c. 400 µm in 

diam., roundish to more often ±irregular or angular, apothecioid, solitary to fused and 

clustered, often forming stroma-like structures, immersed. Disc predominantly entirely visible 

from surface, grayish, pruinose. Pores wide, up to c. 350 µm in diam., formed by proper 

exciple, proper exciple apically or in upper parts becoming visible from surface, roundish to 

±irregular or angular, predominantly entire, free, short, whitish to off-white, often ±shrunken, 

erect to slightly recurved. Thalline rim margin gaping, moderately thin to moderately thick, 

roundish to ±irregular or angular, entire to slightly eroded, brighter than thallus or 

concolorous with thallus. Proper exciple becoming partly to entirely free, moderately thin, 

hyaline to pale yellowish internally, orange to pale brownish marginally, non-amyloid, often 

with old and deceased ascospores in transitional zone between hymenium and proper exciple. 

Hymenium up to c. 120 µm high, non-inspersed, moderately conglutinated, paraphyses 

straight to slightly bent, interwoven, slightly branched towards the margins and the apical 

hymenium, with distinctly thickened tips, lateral paraphyses and true columella absent, in 

fused ascomata sometimes columella-like structures present. Epihymenium moderately thick, 

hyaline, with grayish to pale brown granules. Asci 8-spored, tholus (moderately) thick, thin 

when mature. Ascospores (very) small, submuriform, cell walls moderately thick, endospore 

thickened, non-halonate, brown, non-amyloid to rarely very faintly amyloid, subglobose to 

oblong to ellipsoid, with roundish to narrowed-roundish ends, loci roundish, subglobose to 

oblong, with same shaped end cells, transverse septae (moderately) thin, regular, 7-17 x 5-9 

µm with 4-6 x 1-3 loci. Pycnidia present, immersed with darkened pore area, conidia variable, 

roundish to oval or fusiform, up to c. 3 x 1 µm. 

CHEMISTRY – Thallus K+ yellowish to brown, C-, PD+ orange; containing constictic, 

stictic, hypostictic (majors), α-acetylconstictic, hypoconstictic and cryptostictic (traces) acids. 


glaucescens was collected in Australia on tree bark 

in (sub)tropical rainforests in altitudes ranging from 

sea level to 900 m. It is rare but wide-spread, 

occurring in northern and southern Queensland. 

This is the first report for Australia. Previously this 

pantropical species has been recorded from the 

Neotropics, India and Sri Lanka (Hale, 1981). 

NOTES – This taxon is characterized by the thick, 

bulging thallus with many crystal in-clusions, the 

immersed, irregular/angular ascomata with free 

exciple that usually are clustered and forming 

stroma-like arrangements, the small, ±roundish, 

submuriform, brown ascospores with distinctly 

thickened parts and the stictic acid chemosyndrome. 

Some specimens, in particular Lumbsch 

& Mangold 19117 g have almost chroodiscoid 

apothecia and thus remind of Reimnitzia santensis 

(see fig. 55, D). Besides the chemical differences 


L. glaucescens. 

(R. santensis lacks secondary substances), the ascospore and ascomata size is different 

(ascomata up to 2 mm in diam., ascospores up to 25 x 12 µm). Leucodecton albidulum is 

another similar species, but differs in having larger, transversely septate ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Crystal Cascades, 5 km W of Cairns, Lumbsch & Mangold 

19117 g (F). Atherton Tablelands: Along west boundary of Lake Eacham NP., Hale 830700, 831003 (US);


Wongabel SF., S of Atherton on the Kennedy Hwy., Hale 832559 (US). Edmund Kennedy NP.: Few km N of 

Cardwell, A. & K. Kalb 34283 (hb. Kalb); On Clift Rd., Hale 832716 (US). Kalpowar Forest Drive. c. 40 km NE 

of Monto, SW of Gladstone, Hale 831385, 831427 (US). Upper Coomera, Wilson s.n. (NSW n. 539384 pr. p.). 

Mt.Mee SF., near Mt.Mee, N of Brisbane, Hale 58568 (US). 

Leucodecton occultum (Eschw.) A. Frisch 

Bibl. Lichenol. 92: 157 (2006). Bas.: Thelotrema occultum Eschw., in Martius, Eschweiler & Nees ab 

Esenbeck: Flora Brasiliensis I, 1: 174 (1833). Type: Brazil, Caitété (Bahia), s.c., pr.p. (M-lectotype, selected by 

Hale [1974a; 40]; G-isolectotype). 

Urceolaria compuncta Ach., Meth. Lich.: 143 (1803) [non Porina compuncta Ach., Syn. Lich.: 112 (1814)]. 

Thelotrema compunctum (Ach.) Nyl., Mém. Soc. Sci. Nat. Cherbourg 5: 118 (1858). Leptotrema compunctum 

(Ach.) Müll. Arg., Flora 71: 527 (1888c). Leptotrema compactum (Ach.) Müll. Arg., Flora 70: 400 (1887b) 

[erroneous]. Myriotrema compunctum (Ach.) Hale, Mycotaxon 11: 133 (1980). Type: Indonesia, Chr. Smith s.n. 

(LINN-Sm. 1692.7-lectotype, selected by Hale [1981: 276]; H-Nyl. 22447!-, G!-isolectotypes). 

Leptotrema compunctum var. purpuratum Müll. Arg., Bull. Soc. Roy. Bot. Belgique 30: 75 (1891d). Type: 

Costa Rica, 1890, Pittier 5218 (G-holotype, US!-isotype). 

Thelotrema loandense Vain., in Hiern, W. P. (edit.): Catalogue of the African plants collected by Dr. 

Friedrich Welwitsch in 1853-1861 2(2): 429 (1901). Leptotrema loandense (Vain.) Zahlbr., Cat. Lich. Univ. II: 

636 (1923). Type: Angola, Mar. 1854, Welwitsch 441 (TUR-Vain. 26793-holotype, BM!-isotype). 

Thelotrema compunctum var. antillarum Vain., Ann. Acad. Sci. Fenn., ser. A 6(7): 134 (1915). Leptotrema 

compunctum var. antillarum (Vain.) Zahlbr., Cat. Lich. Univ. II: 633 (1923). Type: Virgin Islands, 14.02.1906, 

Raunkiaer 553 (TUR-Vain. 26798-lectotype, selected by Salisbury [1971a: 276]; C!-isolectotype). 

Thelotrema compunctum var. praiense Vain., Boletim Soc. Broteriana, sér. 2, 6: 149 (1929). Type: 

Mozambique, 1917, Pires de Lima 383 (TUR-Vain. 34802 lectotype, selected by Salisbury [1971a: 276]). 

Thelotrema compunctum f. portoricensis Vain., Mycologia 21: 38 (1929). Leptotrema compunctum f. 

portoricense (Vain.) Zahlbr., Catal. Lich. Univ. II: 632 (1923). Type: Porto Rico, 01.03.1916, Fink 1749 (Wholotype, 

NY!-isotype). 


Thallus epi- to hypophloedal, variable in thickness, thin to moderately thick, up to c. 500 

µm high, pale grayish to pale greenish- or yellowish-gray. Surface dull to slightly shiny or 

somewhat glittering, smooth to more often ±roughened to porous, continuous to rugose to 

rarely somewhat verrucose, distinctly fissured. True cortex absent, cortex-structures absent or 

rarely covered by an incontinuous protocortex up to c. 10 µm thick. Algal layer poorly 

developed, incontinuous, calcium oxalate crystals usually abundant, small to more rarely 

large, scattered to clustered. Vegetative propagules not seen. Ascomata moderately 

conspicuous, small to moderately large, up to c. 600 µm in diam., roundish, apothecioid, 

solitary, immersed to slightly emergent, then hemispherical with same surface as thallus. Disc 

partly visible from surface, grayish, coarsely pruinose. Pores small to moderately wide, up to 

c. 200 µm in diam., predominantly formed by proper exciple, proper exciple usually apically 

or in upper parts becoming visible from surface, roundish to irregular with entire to 

indistinctly split or slightly dentate margin, whitish, sometimes shrunken, incurved to erect to 

rarely somewhat recurved. Thalline rim margin wide to gaping, moderately thin to moderately 

thick, roundish to slightly elongate or irregular, entire, incurved, thalline rim concolorous 

with thallus, sometimes circularly fissured along the ascoma outline, forming a ring-like 

structure. Proper exciple free in the upper parts, moderately thin to moderately thick, hyaline 

to pale yellowish or pale brownish internally, yellowish- to reddish-brown or brownish 

marginally, sometimes slightly carbonized or dark-brown apically, non-amyloid. Hymenium 

up to c. 170 µm high, non-inspersed, strongly conglutinated, paraphyses ±straight, somewhat 

irregular and with ±distinct septation, distinctly interwoven, parallel to slightly branched, 

particularly towards the margins and the epihymenium, with slightly thickened, sometimes 

±irregular tips, lateral paraphyses and columellar structures absent. Epihymenium moderately


thick, hyaline, with grayish-brown granules and small crystals. Asci 8-spored, distinct tholus 

usually not developed, in younger asci with moderately thickened walls throughout, at 

maturity distinctly thinning only in lower parts. Ascospores (moderately) small, (sub- 

)muriform, cell walls thick, endospore (moderately) thin, non-halonate, becoming distinctly 

brown at early maturity, non-amyloid to faintly amyloid only in younger ascospores, oblong 

to predominantly ellipsoid with rounded to narrowed-rounded to rarely slightly subacute ends, 

loci roundish to slightly angular, predominantly subglobular to somewhat irregular, with same 

shaped end cells, transverse septae moderately thin to moderately thick, predominantly 

regular, 20-35(40) x 10-17 µm with 6-8(10) x 1-5(6) loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ orange-red, C-, PD-; containing norstictic (major), stictic (minor 

to absent), α-acetylconstictic (minor to absent) and connorstictic (minor to absent) acids. 

Fig. 56. Leucodecton occultum: growth habit (A), ascomata (B), younger ascospore (C) and older 

ascospore (D). A.: NY-isotype of T. compunctum f. portoricensis; B.-D.: Streimann 48587. Bar= 

A: 2 mm; B: 1.25 mm; C: 5 µm; D: 6 µm. 

ECOLOGY AND DISTRIBUTION – Leucodecton occultum occurs on tree bark in tropical 

mangroves, more rarely in (sub)tropical rainforests in altitudes ranging from sea level to 900 

m. It is common and wide-spread in the north-western Northern Territory and Queensland. 

The pantropical species is also known from the Neotropics, Africa, India (Sethy & al., 1987), 

Sri Lanka (Hale, 1981), Philippines (ibd.) and New Caledonia.


NOTES – It is a common taxon that is characterized 

by the ecorticate, fissured thallus, immersed to 

slightly emergent ascomata with free proper 

exciple, small, (sub-)muriform, brown, thick-walled 

ascospores and the norstictic acid chemosyndrome. 

It is similar to L. subcompuntum, which lacks 

norstictic acid, has more distinctly developed cortex 

structures and has slightly larger ascospores (up to 

45 x 20 µm in size). 

SPECIMENS EXAMINED – Australia, Northern Territory: 

Darwin area, Stevens 4171, 4173 (BRI). Darwin and Gulf 

District, Thompson 492 (CANB). 8 km E of Darwin, N of 

Landing Quarantine station, Thor 5918, 5927 (S). Meckitt 

creek landing, 18 km NE of Darwin, Thor 5828 (S). Kakadu 

NP., A. & K. Kalb 29583, 30472, 30509, 30576, 30593, 

30596, 30778 (hb. Kalb). Daly River Rd. at Adelaide River 

Crossing, 27 km S of Adelaide River Settlement, Lumbsch & 

Elix 8841 (F). Greenant Creek, 35 km SSE of Adelaide River 

Settlement, Streimann 48587 (B). Stuart Hwy., 11 km NW of 

Pine Creek, Streimann 42110 (CANB). 116 km SSW of 

Darwin, Adelaide river near the road, Thor 5995d (S). Green 

Ant Creek, 35 km SSE of Adelaide River Settlement, Elix 28188, Streimann 48587 (CANB). Queensland: 

Uncertain locality, Knight s.n. (G-10194/8). Cape York Peninsula: Cook, Pennefather River, Morton 1033305 

(MEL); Weipa, Specht 689029 (BRI); At Musgrave, on Peninsula Developmental Rd., Hale 831171 (US). Mt. 

Lewis Road, W of Mossman, Hale 831932 (US). Cairns, 5 km N of the city near the airport, A. & M. Aptroot 

22170, 22191 (ABL). Conway Range NP., near Shute Harbour-Airlie Beach, Hale 831738, 830884 (US). Mt. 

Archer Environmental Park, 7 km NE of Rockhampton, Elix 34542 (CANB). Ross Creek, Yeppoon, Elix 34599, 

34619 (CANB, B). Uncertain locality, 'Bailey Book', p. 2, s.c. (BRI-AQ720155). 

Leucodecton subcompunctum (Nyl.) A. Frisch 

Bibl. Lichenol. 92: 162 (2006). Bas.: Thelotrema subcompunctum Nyl., Bull. Soc. Linn. Normandie, sér. 2, 

2: 76 (1868). Leptotrema subcompunctum (Nyl.) Zahlbr., Cat. Lich. Univ. II: 640 (1923). Myriotrema 

subcompunctum (Nyl.) Hale, Mycotaxon 11: 135 (1980). Type: New Caledonia, Lifu, Loyalty, 1864, Thiébaut 

s.n. (H-Nyl.22440!-lectotype, selected by Hale [1974b: 42]; G!-, PC-isolectotypes). 

Leptotrema diffractum Müll. Arg., Hedwigia 30: 50 (1891c). Type: Australia, Queensland, Bellenden Ker, 

Bailey 527 (G!-holotype, BRI!-isotype). 

Leptotrema polycarpum Müll.Arg., Bull. Herb. Boissier 3: 315 (1895). Type: Australia, Queensland, 1887, 

Knight 80 (G!-holotype, G!-isolectotypes [6 samples]). 

Leptotrema inclusum Zahlbr., Bot. Mag. Tokyo 41: 317 (1927). Type: Japan, Kinkuwasan, June 1902, Faurie 

5147 (W-holotype). 

Myriotrema decorticatum Hale, Bull. Br. Mus. Nat. Hist. (Bot.) 8(3): 277 (1981). Type: Sri Lanka, Matara 

District, Hale 46259 (US!-holotype, AMH-, PDA-isotypes). 



L. occultum. 

Thallus epi- to hypophloedal, moderately thin to moderately thick, up to c. 400 µm high, 

variably colored, pale grayish- to yellowish-green or (pale) olive. Surface variable, dull to 

somewhat shiny, smooth to roughened or porous, continuous to verrucose or verruculose, 

unfissured to distinctly fissured to areolate. Thalline cover variable, in younger specimens 

thallus sometimes without cortex structures, otherwise covered by a ±continuous protocortex 

up to c. 30 µm high, sometimes becoming conglutinated, forming a true cortex of irregular to


Fig. 58. Leucodecton subcompunctum: growth habit and ascomata (A-C), ascoma section (D) and 

young, mature and over-mature ascospores (E-G). A.: Lumbsch & Mangold 19116 o; B.: Hale 

830876; C., F.: H-lectotype; D., E.: US-holotype of M. decorticatum; G.: G-holotype of L. 

diffractum. Bar= A: 1 mm; B: 1.5 mm; C: 0.8 mm; D: 100 µm; E: 12 µm; F, G: 20 µm. 

rarely periclinal hyphae. Algal layer well developed and continuous, calcium oxalate crystals 

small to large, scattered to clustered, often forming a continuous layer in the lower medulla. 

Vegetative propagules not seen. Ascomata inconspicuous to moderately conspicuous, 

(moderately) small, up to c. 400 µm in diam., predominantly roundish to slightly irregular, 

becoming apothecioid, solitary to marginally fused, sometimes clustered, predominantly 

immersed to slightly raised to rarely distinctly emergent, then hemispherical to slightly 

subglobose and with same surface as thallus. Disc often becoming partly visible from surface, 

grayish, slightly to distinctly pruinose. Pores small to moderately wide, up to c. 300 µm in 

diam., predominantly formed by proper exciple, proper exciple apically or in upper parts 

usually becoming visible from surface, becoming predominantly free in mature stages, short, 

whitish to grayish or brownish, sometimes shrunken, incurved to erect to rarely somewhat 

recurved. Thalline rim margin wide to gaping, moderately thin to moderately thick, roundish 

to slightly elongate or irregular, entire, incurved, thalline rim concolorous with thallus or 

somewhat brighter. Proper exciple usually becoming free apically or in upper parts, 

moderately thin to moderately thick, hyaline to pale yellowish or pale brownish internally, 

yellowish- to grayish-brown or brownish marginally, sometimes slightly carbonized or dark-


brown apically, non-amyloid to faintly amyloid at the base. Hymenium up to c. 150 µm high, 

non-inspersed, strongly conglutinated, paraphyses ±bent to often distinctly curly in apical 

parts, distinctly interwoven, unbranched to slightly branched towards the margins, with 

slightly thickened tips, lateral paraphyses and columellar structures absent. Epihymenium thin 

to moderately thick, hyaline, with grayish-brown granules and small crystals. Asci 8-spored, 

tholus (moderately) thin, not visible at maturity. Ascospores variable, very to moderately 

small, (sub-)muriform, cell walls (moderately) thick, endospore moderately thin to 

moderately thick, non-halonate, becoming brown usually at late stage of maturity, nonamyloid 

to faintly amyloid in younger ascospores, oblong to ellipsoid to fusiform, with 

rounded to subacute ends, loci roundish to slightly angular, subglobular to oblong to 

somewhat irregular, with same shaped end cells, transverse septae moderately thin to 

moderately thick, predominantly regular, 10-45(50) x 7-20 µm with (4)6-10(12) x 1-6 loci. 

Pycnidia not seen, fide Frisch (2006: 164) immersed with blackish and sometimes detached 

pore area, conidia irregular, narrow elliptical to obovate, up to 5 x 1.2 µm. 




subcompunctum was collected in Australia on tree 

bark in (sub)tropical to warm-temperate wet 

sclerophyll forests and rainforests in an altitude 

ranging from sea level to 1500 m. It is common and 

wide-spread from northern Queensland to central 

New South Wales and also occurs in Lord Howe 

Island. The pantropical species is also known from 

North America (Harris, 1995), Africa (Frisch, 

2006), Sri Lanka (Hale, 1981), Japan (Zahlbruckner, 

1927) and New Caledonia. Hale (1974b) 

reports L. subcompunctum from Dominica and 

describes the species with 1-2 spores per ascus and 

ascospores up to 55 µm long, which indicates that 

this collections probably belongs to L. compunctellum. 

NOTES – It is a highly variable species, however, 

in the examined material all of the morphotypes are 

linked by intermediate stages. It is characterized by 


L. subcompunctum. 

the absence of a distinct true cortex, the numerous, small, predominantly immersed ascomata 

with partly to entirely free exciples, the moderately small, muriform, brown, thick-walled 

ascospores and the stictic acid chemosyndrome. Myriotrema phaeosporum is similar, but 

distinguished by a less distinct, weakly fissured thallus with a speckled surface pattern and the 

narrow-pored, perithecioid mature ascomata with a fused to indistinctly free proper exciple of 

the Myriotrema-type. For differences to L. occultum, see under that species. Leucodecton 

compunctellum differs by larger ascospores (up to 140 µm long) in1-4-spored asci. 

Myriotrema desquamans and M. trypaneoides have a shiny, distinctly corticate thallus and 

ascoma with smaller pores and fused proper exciple. Leucodecton oxysporum 9 from Brazil is 

another similar species that can be distinguished by ascomata with wider pores, a 

predominantly fused exciple and an inspersed hymenium. Leucodecton fissurinum known 

9 Leucodecton oxysporum (Redgr.) Mangold comb. nov. ined., Bas.: Leptotrema oxysporum Redgr., Hedwigia 

73: 63 (1933). Type: Brazil, Taperinha, Santarem, Jul. 1927, Ginzberger s.n. (B!-lectotype, here selected).


from Sri Lanka and Africa is another close taxon and probably conspecific with L. 

subcompunctum (Hale, 1981: 279f.; Frisch, 2006: 156f). Unfortunately no type material was 

available for this study. However, a collection determined by Hale as ‘Myriotrema’ 

fissurinum from the type locality (Hale 46221) belongs to L. subcompunctum. 

SPECIMENS EXAMINED – Australia, Queensland: Iron Range NP., 11 km SW of Cape Weymouth, Streimann 

56377 (B, CANB). Cape Tribulation area, track to Cape Tribulation Beach, Mangold 32 i (F). Km 45 on 

Mt.Windsor Rd., NW of Mossman, Hale 830759, 831481 (US). Mt. Windsor, NW of Mossman, Hale 830281, 

830843, 830989, 831879, 832062, 832204, 832257, 832330, 832695 (US). Mt.Lewis Rd., W of Mossman, Hale 

830289, 831022, 832779 (US). Near end of Black Mountain Rd., 33 km WNW of Kuranda, Hale 832766 (US). 

End of Clohesy River Rd., W of Cairns, Hale 831053 (US). Atherton Tablelands: Davies Creek Rd., E of 

Mareeba, Hale 830973, 830991, 832623 (US); Danbullah Drive, E of Tinaroo, Hale 832484 (US); Lake Tinaroo, 

Downfall Creek Camping Area, Lumbsch & Mangold 19125 i (F); Lake Euranoo, Lumbsch & Mangold 19127 e, 

f, zd (F); Just S of Tolga on Kennedy Hwy., Hale 831423, 832478 (US); Lake Barrine NP., Hale 830944, 

832415 (US), Filson 515536 (MEL); Lake Eacham NP., Hale 830904, 832730 (US), Mangold 29 ai, p, q, r, u, 

al, bb (F); Plath Rd. logging head, S of Atherton, Hale 831019, 832308, 832453 (US); Palmerston NP., Hale 

831907, 832494, 832522 (US); Souita Falls, Lumbsch & Mangold 19153 p (F); Tumoulin Rd., 5 km from 

turnoff to Ravenshoe, Mangold 30 p, zf, zr (F); 13 km S of Ravenshoe on Tully Falls Rd., Hale 832141 (US). 

Bellenden Ker: Bailey s.n. (NSW-539376); 7 km W of Bellenden Ker, A. & M. Aptroot 22485 (ABL). Francis 

Range, NW of Innisfail Hale 831021, 832114 (US). Culpa logging area, SE of Tully Falls Hale 830850, 831923, 

832662, 832713 (US). Dawson logging area, WSW of Tully, Hale 830290, 831002, 831203 (US). State Forest 

area on Tully Rd., S of Mission Beach, Hale 831355 (US). 7.5 km E of Wallaman Falls, W of Ingham, Hale 

830982, 831020 (US). About 7.5 km E of Wallaman Falls, W of Ingham, Hale 832675, 832328 (US). Paluma 

Range, between Townsville und Ingham, A. & K. Kalb 25781, 25788 (hb.Kalb). Mt. Spec NP., WNW of 

Townsville, Hale 831873, 832205, 832640 (US). Cape Hillsborough NP., NW of Mackay, Hale 830259, 831719 

(US). Eungella NP.: Near Peases Lookout, Hale 831407 (US); 2 km S of Eungella, Tibell 14711/II (UPS); Finch 

Hatton Gorge, Lumbsch & Mangold 19116 o (F). Waterpark State Forest Park, N of Yeppoon, Hale 69146, 

831441 (US). Kalpowar Forest Drive., SW of Gladstone Hale 831437 (US). Kroombit SF., 53 km E of Biloela, 

Elix 34733 (B, CANB). Bunya Mnts. NP., Hale 831504 (US). Noosa NP.: Palm Grove Track, Hale 831295, 

831297, 831298, 831334, 831417, 831447, 831453, 831497, 831606, 831661, 831672, 831689 (US); Near 

lookout, Thor 4902/I (S); Tanglewood Track, Elix 10366 (CANB). Wooroi State Forest Park, W of Teewantin, 

Hale 830275 (US). Kenilworth SF., SW of Kenilworth, Hale 830758 (US). Upper Coomera, Wilson s.n. (NSW- 

603826). Lamington NP., Hale 830876, 831203, 831572 (US). New South Wales: McPherson Range, Border 

Ranges NP., NE of Wiangaree, Hafellner 16644 (GZU). NW part of Wiangaree Forest Drive, N of Kyogle, Hale 

832556 (US). South Toonumbar SF., 22 km WNW of Kyogle, Streimann 60813A (CANB, B). Whian Whian 

SF., W of Mullumbimby, Hale 831642 (US). Nightcap Forest Drive, N of Lismore, Hale 830968, 832425 (US). 

Nightcap NP., Mnt. Nardi/Mnt. Matheson Track, Mangold 22 zl (F). Dorrigo SF., near Dorrigo, Hale 58740 

(US). Dorrigo NP.: Never Never Picnic Area and Rosewood Creek Track, Mangold 24 a (F); Sassafras Creek 

Track, Mangold 25 r (F). Styx River SF., 68 km E of Armidale, Kalb & Williams 20602 (hb. Kalb). Doyles 

River SF., 95 km SE Walcha, Hale 58634 (US). Chicester SF., 21 km SW of Gloucester, Elix 24993 (CANB). 

Barrington Tops NP., Lumbsch & Filson 8668 c (CANB, F). Wallingat SF., 16 km SW of Forster: Elix 42631 

(CANB); Streimann 44242 a (B, CANB). Katoomba: Kalb 20597 (hb. Kalb); Sep.1889, Wilson '3' (G, H). Lord 

Howe Island: Track from Smoking Tree Ridge to Rocky Run, Elix 42458 (CANB); Soldiers Creek, Elix 42403 

(CANB). Uncertain locality: 'Shirley Book', p. 23, s.c. (BRI-AQ721253). Sri Lanka, Hale 46221, 46303 (US). 

2. 9. 6. Melanotopelia Lumbsch & Mangold, gen. nov. ined. Type species: Topeliopsis 

toensbergii Vezda & Kantvilas [=M. toensbergii (Vezda & Kantvilas) Lumbsch & 

Mangold comb. nov. ined.]. Type: U.S.A., Washington, Tønsberg 25545 (BG!-holotype). 

Melanotopelia rugosa is the only known species in Australia, for a description see there. 

NOTES – This new genus includes two species formerly placed in Topeliopsis. They are 

distinguished from the latter by a distinctly dark to carbonized marginal proper exciple and 

non-amyloid ascospores (a few species in Topeliopsis s. lat. have I- ascospores), which was 

confirmed by molecular data (part 3). A similar exciple structure, viz. dark to carbonized 

marginally, hyaline with lateral paraphyses internally, can be found in Diploschistes (readily 

distinguished by a different photobiont and ecology) and in the ‘Leptotrema schizoloma-


group’. The members of the latter group, however, are distinguished by regenerating 

ascomata with a ±distinctly layered margin. 


Melanotopelia rugosa (Kantvilas & Vezda) Lumbsch & Mangold comb. nov. ined. 

Bas.: Topeliopsis rugosa Kantvilas & Vezda, Lichenologist 32: 351 (2000). Type: Tasmania, Reservoir 

lakes, Kantvilas 177/86 (HO!-holotype, hb. Vezda-isotype). 


Fig. 60. Melanotopelia rugosa: growth habit (A), ascomata (B), ascoma section (C), asci (D), 

ascospore (E) and ascospore detail (F). A.-F.: HO-holotype. Bar= A: 1 mm; B: 0.6 mm; C: 200 

µm; D: 30 µm; E: 25 µm; F: 12.5 µm.


Thallus muscicolous to partly corticolous, epi- to hyposubstratic, thin, up to c. 80 µm high, 

pale off-white. Surface dull to shiny or glittering, smooth, continuous, unfissured. True cortex 

present, continuous to incontinuous, formed of periclinal hyphae, up to c. 30 µm thick. Algae 

layer poorly developed, incontinuous, calcium oxalate crystals absent. Vegetative propagules 

not seen. Ascomata typical, conspicuous, (moderately) large, up to c. 1 mm in diam., 

roundish, perithecioid first, becoming apothecioid with maturity, sessile, solitary to 

marginally slightly fused, often growing successively, distinctly emergent, subglobular in 

younger stages, becoming urceolate. Disc often becoming partly visible from surface, whitish 

to pale brownish, epruinose, often distinctly cracked or eroded with age. Proper exciple not 

visible from surface, pores small to gaping, up to c. 600 µm in diam., in younger ascomata 

irregular star-shaped, becoming roundish in older ascomata, pore margin entire to slightly 

split. Thalline rim in young ascomata concentrically rugose at the apex, whitish to pale 

grayish, in older ascomata thalline rim margin only slightly rugose to finally entire, remaining 

whitish to pale grayish, forming a conspicuous contrast to the otherwise dark-gray to blackish 

thalline rim, incurved. Proper exciple fused, thick, hyaline internally, dark-brown to 

carbonized marginally, often amyloid towards the base and the subhymenium. Hymenium up 

to c. 350 µm high, non-inspersed, moderately conglutinated, paraphyses straight, distinctly 

parallel, unbranched, with slightly thickened tips, lateral paraphyses present, conspicuous, not 

clearly separated from proper exciple, up to c. 50 µm long, columellar structures lacking. 

Epihymenium indistinct and hyaline in younger ascomata, becoming (moderately) thick, 

reddish to dark brown with age, granules or crystals absent. Asci 1-4-spored, tholus 

(moderately) thick, thin when mature. Ascospores moderately to very large, densely 

eumuriform, cell walls distinctly thickened in younger ascospores, becoming thin in later 

stages, endospore (moderately) thin, sometimes with thin halo, hyaline, yellowish to pale 

brownish in old or decayed ascospores, non-amyloid to very faintly amyloid, predominantly 

fusiform to irregular-fusiform with narrowed-roundish ends, loci roundish to somewhat 

angular, predominantly irregular, transverse septae distinct and regular in younger ascospores, 

becoming indistinct or lacking at late maturity, 80-230 x 20-45 µm with multiple loci. 

Pycnidia not seen. 


stictic (majors), and cryptostictic (minor) acids. [Fide Kantvilas & Vezda (2000) some 

specimen lacking secondary metabolites.] 

ECOLOGY AND DISTRIBUTION – Melanotopelia 

rugosa was collected in Australia predominantly on 

bryophytes in subalpine moorland in altitudes 

ranging from 750-1050 m. One specimen was 

collected on tree bark in a tropical rainforest at 

1500 m. It is a rare, disjunct species, occurring in 

south-west Tasmania and in high altitudes in 

Queensland. It is also known from New Zealand 

and subantarctic islands (Lumbsch & al., 2008), 

indicating a subantarctic distribution extending to 

high altitudes in the tropics. 

NOTES – This taxon can be easily recognized by 

the thin, corticate thallus, the urceolate ascomata 

with bright margins and a dark thalline rim, a 

distinctly carbonized exciple, large, hyaline, 

densely eumuriform ascospores and the stictic acid 


M. rugosa.


chemosydrome. The only Australian species, which could be confused with M. rugosa, is T. 

muscigena with similar ascospores that differ in being strongly amyloid. Moreover, it 

contains no secondary compounds and lacks carbonization. Melanotopelia toensbergii from 

north-Pacific U.S.A., the only other thus far known member of Melanotopelia, differs in 

having smaller ascomata with smaller pores, smaller ascospores (up to 110 µm long) and 

containing the protocetraric acid chemosyndrome. 

SPECIMENS EXAMINED – Australia, Queensland: Bellenden Ker NP., centre peak, 40 km S of Cairns, A. & M. 

Aptroot 22535 (ABL). 

2. 9. 7. Myriotrema Fée, Méthode lichénographique et genera. Paris: 34 (1824). Type species: 

Myriotrema olivaceum Fée. 

Coscinedia Massal., Atti Reale Ist. Veneto Sci. Lett. Arti, ser. 3, 5: 256 (1860). Type species: Coscinedia 

micropora (Mont.) Massal. Type: Indonesia, Java, Junghuhn 143 (PC-holotype). 

Ocellis Clements, The genera of fungi: 80 (1909). Type species: Ocellis myriopora (Tuck.) Clements. Type: 

Cuba, Monte Verde, Wright 129 (FH-Tuck.-holotype, H-Nyl. 22646-isotype). 

THALLUS – Crustose, rarely bulging and flaking away from substrate, corticolous, thin to 

thick, epi- to hypophloedal, up to c. 100-1000 µm high, mostly in shades of olive or gray, 

with greenish to yellowish or whitish tones, sometimes with pale patches due to a partly 

absent algal layer or with a grainy-speckled pattern. Surface shiny to dull, smooth, usually 

continuous to rugose or slightly verrucose, rarely verrucose or verruculose, unfissured to 

fissured or rarely distinctly areolate or rimose. Prothallus thin to indistinct, pale to darkish 

brown. Corticate to rarely ecorticate, either with a ±distinct, continuous true cortex at least in 

younger thalli, hyaline to slightly yellowish, up to c. 20-60 µm thick, sometimes with distinct 

internal splitting, consisting of periclinal to more rarely irregular hyphae; or covered by a 

continuous to discontinuous protocortex up to c. 30 µm thick. Algal layer well-developed, 

rarely poorly developed, continuous to discontinuous, calcium oxalate crystals abundant to 

lacking, small to more often large, scattered to more often clustered, sometimes forming 

columnar structures. Often with distinct medulla layer. Vegetative propagules present or 

absent, isidia occurring in two Australian species (M. eminens, M. frustillatum) for 

description see under these species. 

ASCOMATA – Predominantly inconspicuous, rarely conspicuous, often small, up to c. 200- 

500 mm in diam., or large, up to c. 600-800 µm in diam., roundish to more rarely ±irregular, 

rarely elongate to distinctly lirelliform. Peri- to apothecioid, solitary to distinctly fused, 

sometimes clustered and forming patches of densely arranged ascomata, rarely forming 

stroma-like structures, non-regenerating, often distinctly immersed to slightly raised, or rarely 

distinctly emergent and sessile, then usually hemispherical to conical, rarely subglobose or 

urceolate. Disc often not visible from surface, sometimes partly visible and flesh colored, 

predominantly epruinose to pruinose. Pores mostly tiny to moderately small, up to c. 30-200 

µm in diam., rarely wide or gaping, up to c. 300-600 µm in diam., roundish to ±irregular, 

rarely elongate. Proper exciple not visible from the surface, pore margin entire to slightly, 

rarely distinctly split, formed by thalline rim; or proper exciple becoming ±visible from 

surface, free to fused, off-white to whitish, entire to split, sometimes shrunken, incurved. 

Thalline rim margin small to wide, thin to thick, entire to slightly split, rarely distinctly split 

and/or lacerate to eroded, concolorous with thallus to more often ±brighter or distinctly 

whitish, off-white or pale brownish, incurved to rarely ±erect. Proper exciple fused to slightly 

detached or distinctly free, thin to moderately thick, hyaline to pale yellowish, rarely orange 

internally, yellowish to reddish or grayish to brownish marginally, apically often darkened or


covered by granules, non-amyloid to amyloid basally. Subhymenium indistinct, thin and 

concolorous with basal exciple. Hymenium non-amyloid, moderately to distinctly cupular, up 

to c. 70-300 µm high, predominantly non-inspersed and clear, rarely inspersed or 

interspersed, usually moderately to strongly, rarely weakly conglutinated. Paraphyses often 

±thickened, irregular and/or distinctly septate, rarely straight to usually ±bent, rarely distinctly 

curly in apical parts, parallel to more often ±distinctly interwoven, often ±distinctly branched, 

in particular towards the margins and tips, tips predominantly ±thickened, often ±irregular. 

Lateral paraphyses and true columella absent, in fused ascomata often columella-like 

structures present. Epihymenium mostly indistinct to thin, more rarely ±thick, hyaline, 

egranulose or granulose. 

Asci 8-spored, rarely 1-2-spored, non-amyloid, clavate, ascus walls mostly not thickened, 

tholus predominantly present, ±thick in younger stages, ±thin to not visible at maturity, in M. 

protoalbum with a distinctly tapered ocular chamber; rarely with moderately thickened ascus 

walls and without distinct tholus. Ascospores uni- to biseriate, small to very large, 10-200 x 

4-50 µm, transversely septate to submuriform or eumuriform. Cell walls predominantly 

±thick, rarely (moderately) thin, smooth, endospore in muriform ascospores thin to thick, 

usually non-halonate to indistinctly halonate, rarely distinctly halonate, hyaline to sometimes 

distinctly brown, amyloid to non-amyloid, rarely subglobose or claviform to often oblong to 

ellipsoid or fusiform, with roundish to subacute, rarely acute ends, with 1-12 x 0-7 or multiple 

loci, loci roundish to slightly, rarely distinctly angular, subglobose to oblong or lentiform, 

more rarely ±irregular, with same shaped, hemispherical or conical end cells, transverse 

septae thin to moderately thick or rarely thick, distinct to sometimes indistinct with age in 

densely muriform ascospores, regular to irregular. 

PYCNIDIA – Often present, immersed or in thallus warts with dark to brownish pore area, 

often surrounded by a brighter zone, conidia bacilliform, fusiform to very rarely irregular, up 

to c. 5-8 x 1-2 µm. 

CHEMISTRY – β -Orcinol depsidones present or absent; M. olivaceum contains various 

hitherto unknown substances (‘olivaceum unknowns’). 

ECOLOGY AND DISTRIBUTION – The Myriotrema species in Australia occur on tree bark, in 

altitudes ranging between sea level and 1500 m. The species were predominantly found in 

subtropical to tropical rainforests, more rarely also in wet sclerophyll forests, mangroves and 

monsoon forests of the (sub)tropics, in north-western Northern Territory, Queensland, 

Norfolk Island, northern New South Wales and Lord Howe Island. At present state of 

knowledge, amongst the 17 species known in Australia, two are endemic (M. frustillatum, M. 

temperatum), seven are paleotropical to paleosubtropical (M. desquamans, M. eminens, M. 

microporum, M. phaeosporum, M. polytretum, M. protoalbum, M. subconforme) and eight 

are pansubtropical to pantropical (M. album, M. clandestinum, M. glaucophaenum, M. 

myrioporum, M. olivaceum, M. rugiferum, M. trypaneoides, M. viridialbum). 

NOTES – This genus was reintroduced by Hale (1980, 1981) to accommodate taxa lacking 

carbonization and lateral paraphyses (Thelotrema sect. Myriotrema in Salisbury [1971a, 

1978]). Frisch (2006) excluded several taxa based on morphological differences and partially 

supported by molecular data, and assigned them to various other genera (e.g. Fibrillithecis, 

Leptotrema, Leucodecton, Ocellularia, Stgeobolus). In its present circumscription the genus is 

still considered heterogeneous, Frisch (ibid.) separates two major groups, the ‘M. olivaceumgroup’ 

and the ‘M. viridialbum-group’. In the present treatment, Frisch’s classification is 

accepted in large parts, however, only the ‘M. olivaceum-group’ is confirmed as a well 

separated sub-group as described by Frisch (ibid.). The distinguishing character of a reticulate


columella in the ‘M. viridialbum-group’, which, according to Frisch, sets the sub-group in 

close affinity to Stegobolus, remains dubious. Columella-like structures are found in many 

species, not only in M. glaucophaenum and M. viridialbum (the two Australian species 

grouped in the ‘M. viridialbum-group’) but also in members that show strong similarities to 

the ‘M. olivaceum-group’ (M. album, M. rugiferum), these structures do not represent a true 

columella as found in e.g. Ocellularia or Stegobolus, but are formed due to the fusion of 

individual ascomata (see also under M. glaucophaenum). Further, several taxa are included in 

Myriotrema which are deviant from both sub-groups, as M. desquamans, M. phaeosporum or 

M. trypaneoides with brown ascospores reminiscent to Leucodeton, or species with large, 

densely muriform ascospores (M. eminens, M. frustillatum). Hence, I accept here a 

Myriotrema s. str. group and Myriotrema s. lat., including all remaining species placed in the 

genus for the time being. 

Myriotrema s. lat. is circumscribed by a ±pale proper exciple consisting of 

prosoplectenchymatous hyphae that are distinctly radiating towards the apical parts 

(‘Ocellularia-type’) without substrate inclusions, absence of lateral paraphyses, peri- to 

apothecioid ascomata and predominantly shiny, ±distinctly corticate thalli. Myriotrema s. str. 

is further characterized by small, immersed to slightly raised ascomata, and small, hyaline, 

amyloid transversely septate to submuriform ascospores. Similar genera include Fibrillithecis, 

Leptotrema, Leucodecton and Ocellularia (not treated here). Except for Ocellularia the 

genera are chiefly distinguished by the structure of the proper exciple (see also under this 

genera). The distinction of Ocellularia is more problematic and in certain cases poorly 

understood. The two genera are distinguished by the presence of a true columella and/or 

distinct carbonization. The placement of several taxa, particularly those with absent columella 

and indistinct carbonization, currently grouped in Ocellularia needs further study. 


Myriotrema album Fée 

Essai Crypt. Écorc.: 104 (1824). Thelotrema album (Fée) Nyl., Ann. Sci. Nat. Bot. 5(7): 315 (1867). 

Ocellularia alba (Fée) Müll. Arg., Mem. Soc. Phys. Geneve 29(8): 6 (1887). Thelotrema myriotrema Nyl., 

Memoir. Soc. Sienc. Nat. Cherbourg 5: 107 (1857) [nom. nov. pro Myriotrema album Fée]. Type: "America 

meridionali, ad corticem Bonplandiae trifoliatae (Wild.)", s.c. (G-Fée 249!-lectotype, selected by Hale [1978: 

12]; H-Nyl. 22635-isolectotype). 


Thallus epi- to hypophloedal, (moderately) thick, up to c. 500 µm high, (pale) olive to 

yellowish-olive, often with paler patches due to a partly absent algal layer. Surface ±shiny, 

smooth, continuous to rugose, unfissured. True cortex usually present, predominantly 

continuous, slightly yellowish, thickness variable, up to c. 40 µm thick, consisting of 

periclinal to irregular hyphae, sometimes partly not conglutinated forming a protocortex. 

Algal layer well developed, continuous but sometimes partly thinning or lacking (see above), 

presence of calcium oxalate crystals somewhat variable, predominantly abundant, small to 

large, clustered, sometimes forming short layers, more rarely sparse, then usually found in 

scattered (large) clusters, distinct medullar layer present. Vegetative propagules not seen. 

Ascomata usually inconspicuous, predominantly ±small, up to c. 350 µm in diam., roundish, 

apothecioid, solitary to marginally to rarely entirely fused, often clustered, forming patches of 

densely arranged ascomata, in some specimen ascomata sometimes becoming stroma-like 

fused, then often sharing the same thalline rim (Fig. 62, B), predominantly immersed to rarely


Fig. 62. Myriotrema album: growth habit (A), ascomata (B), ascomata section (C), pycnidia (D), 

conidia (E) and ascospores (F). A., F.: Lumbsch & Mangold 19110 e; B., C.: Streimann 45850; D., 

E.: Mangold 30 r. Bar= A, B: 1 mm; C, D: 100 µm; E: 5 µm; F: 7 µm.


somewhat emergent, then hemispherical. Disc often becoming partly visible from surface, 

pale flesh colored, epruinose to slightly pruinose. Pores (moderately) small to sometimes 

becoming wide, up to c. 200 µm in diam., predominantly roundish to somewhat irregular, 

entire to rarely slightly split, proper exciple usually not visible from surface, in distinctly 

fused ascomata (see above) becoming visible from surface, entire, erect, off-white. Thalline 

rim margin variably thick, usually ±thin to more rarely moderately thick, brighter than thallus, 

forming a narrow to moderately wide off-white ring-like area, level with thallus to often 

somewhat raised, rarely sunken, thalline rim predominantly incurved. Proper exciple fused, 

predominantly moderately thin, hyaline internally, pale yellowish to yellowish-brown 

marginally, non-amyloid. Hymenium up to c. (80)100 µm high, non-inspersed, strongly 

conglutinated, paraphyses straight to slightly branched, distinctly septate, slightly interwoven, 

with distinctly thickened, somewhat irregular tips, lateral paraphyses and true columella 

lacking, in fused ascomata columella-like structures present (see above). Epihymenium 

indistinct to thin, hyaline, sometimes with fine grayish granules. Asci 8-spored, tholus 

(moderately) thick, becoming moderately thin at maturity. Ascospores (very) small, 

transversely septate, often with a single longitudinal septum, cell walls (moderately) thick, 

endospore moderately thin to moderately thick, sometimes with thin halo, hyaline, 

predominantly distinctly to strongly amyloid, rarely with faint amyloid reaction, ellipsoid to 

fusi- or claviform, predominantly with narrowed-roundish or subacute to more rarely acute 

ends, loci roundish to somewhat acute, subglobose to lentiform or irregular, with same shaped 

to hemispherical or conical end cells, septae thin to moderately thick, often slightly to 

distinctly irregular, 10-25(28) x 4-9 µm with 3-8 (x2) loci. Pycnidia present, in thallus warts 

with brownish pore area surrounded by a bright zone, conidia bacilliform, up to c. 8 x 1 µm. 

CHEMISTRY – Thallus K-, C-, PD-; no secondary compounds detectable by TLC. 

ECOLOGY AND DISTRIBUTION – Myriotrema 

album was collected in Australia on tree bark 

predominantly in (sub-)tropical rainforests in 

altitudes ranging from 10 - 1230 m. It is common 

and wide-spread from northern Queensland to 

northern New South Wales. Besides Australia it 

was reported from the Neotropics, Philippines and 

Solomon Islands indicating a pan(sub)tropical 

distribution. 


unfissured, shiny, corticate thallus, small, 

predominantly immersed ascomata with fused 

proper exciple, transversely septate to indistinctly 

submuriform, hyaline, amyloid ascospores with 

thickened parts and the absence of secondary 

compounds. A morphologically similar species is 

M. clandestinum, which can be distinguished by 

less abundant calcium oxalate crystals and the 

presence of the psoromic acid chemosyndrome. 


M. album. 

Other similar Australian species with the same chemistry include M. protoalbum and M. 

myrioporum, for differences see under these species. 

SPECIMENS EXAMINED – Australia, Queensland: Uncertain locality: 'Peachey's Shrub', Aug. 1889, Wilson s.n. 

(NSW 539375). Iron Range NP., 29 km from western boundary on track to Portland Rds., Hale 830043, 832671 

(US). Cape Tribulation area: Myall Beach, Lumbsch & Mangold 19160 w (F); 2 km W of main rd. between Oil


Palms and Coopers Creek, Hale 830043, 832671 (US); Milky Pine Rd., 3 km NE of Daintree River Crossing, 

Hale 832218 (US). Thornton Range: At tower turnoff on CREB rd. (to Cooktown), c.15 km N of the Daintree 

River crossing, Hale, 830964, 831194, 831359, 831501, 832558 (US); CREB rd., about 5 km in from Daintree 

River crossing, Hale 831472, 831655 (US); CREB track, Stevens 19184 (GZU). Mt. Windsor logging area, NW 

of Mossman: 9 km from rd. to old Forestry Camp and the main rd., Hale 832542; E of old Forestry Camp, Hale 

832391 (US). Mt. Lewis Rd., 10.5 - 16 km N from Kennedy Hwy., W of Mossman, Hale 69153, 831202, 

832086 , 832337, 832342 (US). Rex Creek, Mossman Gorge, Daintree River NP., 6 km W of Mossman, 

Streimann 45850 (B, CANB). 14 km SW of Mossman, Mt. Lewis, Tibell 14615 (UPS). Cairns, Aug. 1893, 

Wilson s.n. (NSW 539368). Atherton Tablelands: Davies Creek Rd. 17 km S of Kennedy Hwy., S of Davies 

Creek Falls NP., E of Mareeba, Hale 830724 (US); Lamb Range, near Mt. Haig, 20 km SE of Mareeba, 

Streimann 57668 (B); Lake Eacham NP., Mangold 29 aj (F); Plath Rd. logging head, 9 km W of Plath Rd., off 

Kennedy Hwy, Herberton range, S of Atherton, Hale 832554 (US); 13 km SW of Atherton, on summit of Great 

Dividing Range, Weber & Moir 352 (C, CANB, COLO, G, H, MEL, NSW, S, US, WIS); Tumoulin Rd., 5 km 

from turnoff to Ravenshoe, Mangold 30 r (F); 10-18 km S of Ravenshoe on Tully Falls Rd., Hale 832065, 

832798, 832805 (US); Culpa logging area, 13 km from Koombooloomba rd. turnoff, SE of Tully Falls, Hale 

830615 (US). Near TELECOM Cable Car bldg., at base of Bellenden Ker (off Bruce Hwy.), Hale 831531 (US) 

Mt.Tyson, 2 km W of Tully, Streimann 45557 (CANB). State Forest area on Tully Rd., 1 km from jct. with S. 

Mission Beach Rd., S of Mission Beach, Hale 831467 (US). Eungella NP.: "Palm Walk", 2.5 km SE of 

Eungella, A. & K. Kalb 34262, 34263, 34267 (hb. Kalb); NP. side rd. nr. Peases Lookout, off Darymple rd., Hale 

831412, 831598 (US); Rosser Rd. entry point off Darymple rd., near Peases Lookout, Hale 832530 (US); Rd. to 

rainforest at NP. margin, Lumbsch & Mangold 19110 e (F). Waterpark Creek State Forest Park in Byfield SF., 

24 km N of Yeppoon, Hale 831741 (US). Kroombit Tops SF.: Munholme Creek, 47 km SW of Calliope, 

Streimann 65062 (B, CANB); Dawes Range, 53 km E of Biloela, Elix 34741, 34754 (B, CANB, H). Kalpowar 

Forest Drive, c.40 km NE of Monto, SW of Gladstone, Hale 831756, 832173 (US). Wooroi State Forest Park, W 

of Teewantin, Hale 832320, 832724 (US). Mapleton Falls, c. 13 km W of Nambour, Hale 832820 (US). Blackall 

Ranges, 06. Aug. 1890, Wilson s.n. (NSW 539373). Mt. Mee SF., 6 km NW of Forestry Office, NW of Mt. Mee, 

Hale 832790 (US). Mt. Glorious, Brisbane SF., c. 40 km WNW von Brisbane, A. & K. Kalb 26573 (hb. Kalb). 

Brisbane, Aug. 1896, Wilson s.n. (NSW 539372). Upper Coomera, 1802, Wilson s.n. (NSW 539371). 

'Lambourine' [Mnt. Tamborine], 1802, Wilson s.n. (NSW 539374). Carabeen Nature Refuge, 45 km E of 

Warwick, Lumbsch & Mangold 19174 f (F). Lamington NP.: Near lost world, A. & M. Aptroot 21900 (ABL); 

O'Reillys Guesthouse, A. & K. Kalb 21542 (hb. Kalb). New South Wales: Lions Tourist Rd. near Queensland 

border, N of Waingaree, Hale 832764 (US). Nightcap NP., Mnt. Nardi/Mnt. Matheson Track, Mangold 22 zd, zf, 

zg, zh (F). Night Cap Forest Drive, W of Mullumbimby: Big Shrub Flora Reserve, Hale 831375 (US); 

Gibbergunyah Roadside Reserve, Whian Whian SF., Hale 831329, 831400, 831478 (US); 1 km W of Minyon 

Falls, N of Lismore, Hale 832405 (US). Wollongbar, Jun. 1894, Wilson s.n. (NSW 539370). Richmond River, 

Hodgkinson s.n. (M 23370). Iluka Nature Reserve, 50 km NE of Grafton: Mangold 23 r (F); Hale 58546 (US). 

Dorrigo SF., near Dorrigo, Hale 58756 (US). Uncertain locality: 'Shirley Book', p. 23, s.c. (BRI-AQ721249). 

Myriotrema clandestinum (Fée) Hale 

Mycotaxon 11: 113 (1980). Bas.: Thelotrema clandestinum Fée, Ess. Crypt. Suppl. 84: 90 (1837). Type: "In 

america ad corticem Cinchonae lancifoliae Mutis", s.c. (G-Fée 244!-lectotype, selected by Hale [1974a: 32]). 

Thelotrema terebratulum Nyl., Ann. Sci. Nat, Bot, sér. 5, 7: 315 (1867a). Ocellularia terebratula (Nyl.) 

Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 29(8): 12 (1887c). Myriotrema terebratulum (Nyl.) Hale, 

Mycotaxon 11: 135 (1980). Type: Colombia ('Nova Granata'), Rio Negro, 1863, Lindig 129 (H-Nyl. 22637lectotype, 

selected by Hale [1978: 33]; FH-Tuck.!-, G!-, M-, PC-isolectotypes). 

Thelotrema clandestinum f. remanens Nyl., Ann. Sci. Nat., Bot., sér. 5, 7: 317 (1867a) Thelotrema remanens 

(Nyl.) Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 29(8): 7 (1887c). Type: Colombia, Monte del Morro, 

1863, Lindig 90 (H-Nyl.-holotype; BM-, G-, PC-isotypes). 


Thallus epi- to hypophloedal, (moderately) thick, up to c. 700 µm high, (pale) olive to 

yellowish-olive, often with paler patches due to a partly absent algal layer. Surface ±shiny, 

smooth, continuous to rugose, unfissured. True cortex usually present, predominantly 

continuous, slightly yellowish, thickness variable, up to c. 50 µm thick, formed of periclinal 

to somewhat irregular hyphae, rarely partly not conglutinated forming a protocortex. Algal 

layer well developed, continuous but often partly thinning or lacking (see above), calcium


Fig. 64. Myriotrema clandestinum: growth habit (A), pycnidia (B), ascomata section (C) and 

ascospores (D). A.: Lumbsch & Mangold 19132 z; B., C.: Lumbsch & Mangold 19132 s; D.: 

Lumbsch & Mangold 19132 v. Bar= A: 1.5 mm; B: 0.25 mm; C: 250 µm; D: 5 µm. 

oxalate crystals sparse or lacking, usually found in scattered (large) clusters, distinct medullar 

layer present. Vegetative propagules not seen. Ascomata usually inconspicuous, ±small, up to 

c. 300 µm in diam., roundish, apothecioid, solitary to marginally fused, often clustered, 

forming patches of densely arranged ascomata, predominantly immersed to rarely somewhat 

emergent, then hemispherical. Disc often becoming partly visible from surface, pale flesh 

colored, epruinose to slightly pruinose. Pores (moderately) small to sometimes becoming 

wide, up to c. 100(200) µm in diam., predominantly roundish to somewhat irregular, entire to 

rarely slightly split, proper exciple not visible from surface. Thalline rim margin variably 

thick, usually ±thin to more rarely moderately thick, brighter than thallus, forming a narrow 

pale brownish to off-white ring-like area, level with thallus to often somewhat raised, thalline 

rim incurved. Proper exciple fused, moderately thin to moderately thick, hyaline internally, 

pale yellowish to yellowish-gray marginally, non-amyloid. Hymenium up to c. 90(100) µm 

high, non-inspersed, strongly conglutinated, paraphyses straight to slightly branched, 

distinctly septate, slightly interwoven, with slightly to distinctly thickened tips, lateral 

paraphyses and columellar structures absent. Epihymenium indistinct to thin, hyaline, rarely 

with fine grayish granules. Asci 8-spored, tholus moderately thick, thin when mature. 

Ascospores (very) small, transversely septate, sometimes with a single longitudinal septum 

(see notes), cell walls (moderately) thick, endospore moderately thin to moderately thick, 

sometimes with thin halo, hyaline, distinctly to strongly amyloid, ellipsoid to fusiform, 

predominantly with narrowed-roundish to subacute ends, loci roundish to somewhat acute, 

subglobose to lentiform or irregular, with same shaped or hemispherical to conical end cells,


septae moderately thin, sometimes slightly irregular, 10-25(27) x 6-8 µm with 3-6(7) (x2) 

loci. Pycnidia present, in thallus warts with brownish pore area surrounded by a bright zone, 

mature conidia not found, fide Frisch (2006) bacilliform, up to c. 8 x 1.5 µm. 

CHEMISTRY – Thallus K+ yellowish, C-, PD+ yellow; containing psoromic (major) and 

subpsoromic (minor to trace) acids. 


clandestinum occurs in Australia on tree bark 

predominantly in tropical rainforests in altitudes 

ranging from 10 - 900 m. It is common in 

Queensland, occurring frequently in the northern 

part, rarely in northern-central or southern 

Queensland. It is a pantropical species being 

reported from the Neotropics (e.g., Hale, 1978b, 

1981; Sipman, 1992a), Africa (Frisch, 2006), India 

(Nagarkar & al., 1988), Sri Lanka (Hale, 1981), 

Andman Islands (Sethy & al., 1987), Taiwan (Hale, 

1981), Borneo (Sipman, 1993), Philippines (Hale, 

1981), New Caledonia (Nylander, 1868) and 

Solomon Islands (Hale, 1981). 

N OTES – This taxon is characterized by an 

unfissured, thick, shiny thallus that is distinctly 

corticate, the immersed, small ascomata with fused 

exciple, small, hyaline, thick-walled, amyloid 

ascospores that are predominantly transversely 


M. clandestinum. 

septate, and the presence of the psoromic acid chemosyndrome. It is similar to M. album that 

mainly differs by the absence of secondary compounds (for further differences see under that 

species). Myriotrema microporum and M. temperatum are also close species, see there for 

differences. Another similar species is M. glaucophaenum, it can be distinguished by a 

thinner thallus, larger, often distinctly emergent ascomata with a free proper exciple and 

slightly smaller ascospores (up to 20 µm long). Myriotrema terebratulum was considered as a 

similar, but distinct species (e.g., Hale, 1974a: 32, 1978: 41; Patwardhan & Kulkarni, 1977: 

12) distinguished by the strictly transversely septate ascospores. However, in the Fée type 

material of M. clandestinum no submuriform ascospores were found. However, several 

Australian collections have ascospores with an additional longitudinal septum. Hence, I 

follow Salisbury's (1978) opinion to regard the presence or absence of a vertical septum as of 

little taxonomic value. Salisbury (loc. cit.) discussed the contradictory statements of Nylander 

(1867: ascospores 'medio divisis') and Müller (1887: 'clare 4-loculares') for this species. 

Hence T. terebratulum is included as a synonym of M. clandestinum here. 

One of Hale’s collections from Atherton Tableland (Hale 831933) differs by the presence 

of 2-0-demethylsubpsormic acid (Elix, pers. comm.) but lacks psoromic and subpsoromic 

acids. It further differs in strongly aggregated and partly fused ascomata, very similar to some 

collections of M. album. The deviant specimen is tentatively included here for its 

chemotaxonomic relationship, until molecular data are available. 

SPECIMENS EXAMINED – Australia, Queensland: Thornton Range: CREB rd. (to Cooktown), about 5 km in 

from Daintree River crossing, NW of Mossman, Hale 831520 (US). Cape Tribulation Area, Cape Kimberley, 

Lumbsch & Mangold 19166 f (F). Km 45 on Mt. Windsor Rd., NW of Mossman, Hale 831314 (US). Mt. Lewis 

Rd. 4-16 km N from Kennedy Hwy., W of Mossman, Hale 830634, 832588 (US). Daintree NP., Mossman 

Gorge Section, near western border of National Park along Mossman creek, Mangold 36 c (F). Near end of


Black Mountain Rd., 33 km WNW of Kuranda, Hale 832686 (US). Kuranda, Wilson s.n. (MEL 26171). End of 

Clohesy River Rd., 16 km SE Kennedy Hwy., W of Cairns, Hale 832349 (US). Atherton Tablelands: Lake 

Barrine NP., Hale 831577 (US); Lake Eacham NP., track around lake, Hale 830267 (US); W side of Lake 

Eacham NP., 3 km in from W boundary on rd. from Yungaburra, Hale 832353 (US); Area below crater, Mt. 

Hypipamee NP., S of Atherton, Hale 832102, 832129 (US); Malanda Falls, Lumbsch & Mangold 19132 s, v, z 

(F); Millaa Millaa falls, Lumbsch & Mangold 19139 o (F); SW of K-1 tree rd. off Palmerston Hwy, 11 km from 

main hwy and 2 km N of S.Johnstone Forestry Camp, SE of Millaa Millaa, Hale 831933 (US); Palmerston NP., 

6 km E of the West Boundary, Hale 831646 (US); Culpa logging area, 13 km from Koombooloomba rd. turnoff, 

SE of Tully Falls, Hale 832787 (US). Francis Range, Woopen Creek Rd., 25 km in from Bruce Hwy., NW of 

Innisfail, Hale 832677 (US). Mt. Chalmynia logging area, 15 km from Bruce Hwy., W of Innisfail, Hale 832258 

(US). Dawson logging area, State Forest Reserve 605, 24 km S of Koombooloomba turnoff, WSW of Tully, 

Hale 832689 (US). Dicks Tableland, "Rain Forest Discovery track" near Eungella, 60 km W of Mackay, K. & A. 

Kalb 107 (CANB). Python Rock Track, Lamington NP., Hale 830867 (US). 

Myriotrema desquamans (Müll.Arg.) Hale 

Mycotaxon 11: 133. 1980. Bas.: Anthracothecium desquamans Müll.Arg., Flora, Jena 71: 48 (1888). 

Leptotrema desquamans (Müll.Arg.) Patw. & Makhija, Bryologist 83: 368 (1980). Type: Australia, Queensland, 

Johnston River [Innisfail], Berthoud s.n. (G!–lectotype, selected by Patwardhan & Makhija [1980: 368]). 

Thelotrema irosinum Vain., Suomal. Tiedeakat. Toim. A, 15: 174 (1921). Leptotrema irosinum (Vain.) 

Zahlbr., Cat. Lich. Univ. II: 635 (1923). Type: Philippines, Irosin, Elmer 14749 (TUR-Vain.-lectotype, selected 

by Hale [1981: 278]; FH-, G-, L-, W-isolectotypes). 


Thallus epiphloedal to partly hypophloedal, (moderately) thick, up to c. 800 µm high, pale 

olive to greenish gray. Surface shiny, smooth, continuous to slightly verrucose, unfissured. 

True cortex present, up to c. 40 µm thick, consisting of periclinal hyphae. Algal layer well 

developed, continuous but interrupted by calcium oxalate crystals, calcium oxalate crystals 

abundant, small to large, scattered or in clusters. Vegetative propagules not seen. Ascomata 

inconspicuous, small, up to c. 400 µm in diam., roundish, perithecioid, solitary, immersed to 

slightly emergent, then (flattened-)hemispherical. Disc not visible from surface. Pores tiny to 

very small, up to c. 30 µm in diam., roundish, entire, proper exciple apically visible from 

surface, forming the pore margin, fused, usually whitish-translucent, incurved, flush with 

thallus or slightly sunken. Thalline rim margin roundish, whitish or brighter than thallus. 

Proper exciple fused, ±thin, hyaline internally, yellowish- to reddish-brown marginally, often 

±amyloid at the base. Hymenium up to c. 300 µm high, non-inspersed, highly conglutinated, 

paraphyses interwoven, slightly branched, tips thickened, lateral paraphyses and columellar 

structures absent. Epihymenium inconspicuous, hyaline, without granules. Asci 8-spored, 

tholus thick, thin when mature. Ascospores small, eumuriform, cell walls thick in younger 

stages becoming moderately thin at maturity, endospore slightly thickened, non-halonate, 

brown, non-amyloid to faintly amyloid, rarely (sub-)globular (morphotype II) to usually 

oblong, ends roundish to narrowed-roundish to rarely somewhat apiculate, loci roundish to 

slightly angular, (irregular-)subglobose to slightly lentiform, transverse septae thin, regular to 

irregular, 20-35 x 8-18µm, with 6-12 x 1-7 loci. Pycnidia not seen. 

CHEMISTRY – Strain I: Thallus K+ yellowish to brown, C-, PD+ orange; containing 

constictic, stictic, (majors), hypoconstictic, cryptostictic and hypostictic (traces) acids. Strain 

II: Thallus K-, C-, PD-; no secondary compounds detectable by TLC. 

ECOLOGY AND DISTRIBUTION – Myriotrema desquamans grows on tree bark in tropical 

rainforests in altitudes ranging from sea level to 800 m. It is moderately common in northern 

Queensland. This paleotropical species was previously reported from India and the Andman


Fig. 66. Myriotrema desquamans: ascomata (A, B), ascoma and thallus section (C) and ascospores (D, 

E). A.: Lumbsch & Mangold 19166 h; B., C.: Barnsley 1668; D.: Lumbsch & Mangold 19155 e; 

E.: Hale 831005. Bar= A: 0.4 mm; B: 0.5 mm; C: 200 µm; D: 7.5 µm; E: 12 µm 

Islands (as L. irosinum, Patwardhan & Kulkarni, 

1977; Sethy & al., 1987), Sri Lanka (Hale, 1981), 

Philippines (as T. irosinum, Vainio, 1921) and 

Sabah (Patwardhan & Kulkarni, 1977). 


shiny, corticate thallus, the immersed to slightly 

emergent perithecia with small pores, the 

moderately small, brown, muriform ascospores. A 

similar species is M. trypaneoides, see there for 

differences. Other Australian species with brown, 

muriform ascospores and the stictic acid 

chemosydrome (L. glaucescens, L. subcompunctum 

and M. phaeosporum) can be distinguished by 

wider pored ascomata and less distinctly shiny 

thallus surfaces. One collection from Cape 

Tribulation (Mangold 32 zb) differs in having 

globose to subglobose ascospores and is included 

only tentatively under this name until further 


M. desquamans.


collections become available. Further, one Hale collection (Hale 830737) lacks secondary 

compounds but otherwise agrees with the stictic acid containing specimens. Thus it is treated 

as a chemotype (strain II) of M. desquamans. 

SPECIMENS EXAMINED – Strain I: Australia, Queensland: Iron Range NP.: 2-14 km from western boundary on 

track to Portland Roads, Hale 830012, 830017, 830018, 830601, 832807, 832809 (US); Mt. Tozer, Clarkson 

2910 (BRI). Gordon Creek, near Lockhart River settlement, Streimann 56537 (B, CANB). ‘Home Rule’ Track to 

Home Rule waterfall, S of Cooktown, Barnsley 1668 (CANB). Near Cedar Bay NP., on road to Cooktown, 

Mangold 34 m, p (F). Cape Tribulation Area: Emmagen Creek, Lumbsch & Guderley 11161 f (F); Cape 

Tribulation NP., Hale 831709 (US); Cape Tribulation Beach, Hale 831975 (US); Myall Beach, Mangold 32 b, zb 

(F); Buchanan Creek Rd. (Cow Bay), Hale 830260, 831234 (US); Cape Kimberley, Lumbsch & Mangold 

19166e, h, o, 19167b, k, q, r (F). Daintree NP., Mossman Sct., Lower Mossman River, Mangold 35 j, 36 e (F); 

Mossman Gorge, Hale 832207, 832259 (US). Machans Beach, N of Cairns, K. & A. Kalb 21199 (hb. Kalb). 

Fitzroy Island, E of Cairns, A. & M. Aptroot 22305 (ABL). Crystal Cascades, WSW of Cairns, Lumbsch & 

Mangold 19120 d, n (F). Atherton Tablelands: 27 km on Mulgrave River Rd., SW of Gordonvale, Hale 830960 

(US); Danbullah Forest Drive, Hale 69170, 841488, 831515 (US); Curtain Fig Tree, Lumbsch & Mangold 1928 

q (F); W side of Lake Eacham NP., Hale 830938 (US); Souita Falls, Lumbsch & Mangold 19155 e, i (F); SW of 

K-1 tree rd. off Palmerston Hwy, Hale 831005, 830865 (US); Just S of Hwy, 23 km E of jct Kennedy Hwy and 

Palmerston Hwy, Hale 831052 (US); Wooroonooran NP., Palmerston Sct., Lumbsch & Guderley 11108 h (F); 

Kearney Falls, Streimann 46560 b (CANB); Josephine Falls, Hale 831736 (US). 5 km NW of Babinda, near 

bridge crossing of the Russell River, Hale 831115 (US). Babinda Boulders: A. & M. Aptroot 22405 (ABL); Hale 

831594 (US); Mangold 39 za (F). 3 km N of Garradunga, Graham Range, Hale 831605 (US). Francis Range, 

Woopen Creek Rd., Hale 830272 (US). 12 km E of Cardstone on Tully River Rd. to Kareeya Power Station, 

Hale 831970 (US). 

Strain II: Australia, Queensland: Picnic area 2 km E of Cardstone on Tully River Rd. to Kareeya Power 

Station, Hale 830737 (US). 

Myriotrema eminens (Hale) Hale 

Mycotaxon 11: 133 (1980). Bas.: Thelotrema eminens Hale, Mycotaxon 3: 177 (1975). Type: Malaysia, 

Pahang, Hale 29947 (US-holotype, not seen). 


Thallus epi- to hypophloedal, thin, up to c. 100 µm high, whitish-grayish to pale yellowishgray. 

Surface ±shiny, smooth, continuous, unfissured to slightly fissured. True cortex usually 

present, ±continuous, up to c. 20 µm thick, moderately dense, formed of periclinal hyphae. 

Algal layer poorly developed but ±continuous, calcium oxalate crystals sparse to absent. 

Isidia often present, concolorous with thallus, cylindrical, slightly branched, up to c. 1.3 mm 

long and up to c. 200 µm thick. Ascomata conspicuous, moderately large, up to c. 800 µm in 

diam., predominantly ±roundish, perithecioid, marginally fused and clustered, forming groups 

of few to several ascomata, distinctly emergent, subglobose to urceolate. Disc not visible 

from surface. Pores small, up to c. 200 µm in diam., roundish to irregular, entire to slightly 

split, apical proper exciple usually not visible from surface, rarely becoming visible if partly 

detached, entire to slightly split, incurved, whitish, pore area sometimes sunken, concolorous 

with thalline rim or somewhat darkened. Thalline rim margin thick, entire to sometimes 

slightly cracked, flattened to sometimes slightly annular, concolorous with thallus or pale 

yellowish. Proper exciple fused, apically sometimes becoming free, (moderately) thick, color 

conspicuous, orange internally, reddish to reddish-brown marginally, non-amyloid. 

Hymenium up to c. 250 µm high, non-inspersed, strongly conglutinated, paraphyses thick, 

parallel, slightly branched towards the exciple, often distinctly curly in apical parts, tips 

thickened, lateral paraphyses and columellar structures absent. Asci 1(-2)-spored, tholus thin, 

not visible at maturity. Ascospores (very) large, densely eumuriform, cell walls and 

endospore thin, non-halonate, hyaline, distinctly to strongly amyloid, oblong to roundishfusiform 

with roundish to narrowed-roundish ends, often somewhat irregular in outline, loci


small, roundish to slightly angular, ±subglobose, transverse septae thin, irregular, vanishing 

with age, 150-200 x 25-50 µm with multiple loci. Pycnidia not seen. 

Fig. 68. Myriotrema eminens: growth habit (A), ascomata and isidia (B), ascospore (C) and ascospore 

detail (D). A.-D.: Hale 831968. Bar= A: 0.8 mm; B: 0.4 mm; C: 27.5 µm; D: 15 µm. 


C-, PD+ orange; containing constictic, stictic, 

(majors), hypostictic (minor), hypoconstictic and 

cryptostictic (traces) acids. 

ECOLOGY AND DISTRIBUTION – The species was 

collected in Australia on tree bark in tropical 

rainforests in altitudes ranging from 530 to 1000 m. 

It is very rare in northern Queensland. This is the 

first report for Australia, it is known from Sri 

Lanka, Malaysia and the Philippines (Hale, 1981). 

NOTES – Myriotrema eminens is characterized 

by the isidiate, corticate thallus, ±urceolate 

ascomata with a conspicuous, reddish exciple, 

large, eumuriform, thin-walled, hyaline, amyloid 

ascospores and the stictic acid chemosyndrome. 

The number of spores per ascus (one to rarely two) 


M. eminens.


found in the Australian collections is somewhat different from Hale’s (1981) description, who 

gave 2-4 spores per ascus. However, previously (Hale, 1975), he mentioned 1-2 spores per 

ascus for that species. The presence of isidia is variable, the specimen from Atherton 

Tablelands is abundantly isidiate, whereas the sample from Mt. Windsor is almost nonisidiate. 

Variation in isidia occurrence has been described earlier for M. eminens (Hale, 1981: 

279). 

SPECIMENS EXAMINED – Australia, Queensland: Mt. Windsor, 5 km W of new Forestry Camp, NW of 

Mossman, Hale 831880 (US). Off Palmerston Hwy, 11 km from main hwy. and 2 km N of S. Johnstone Forestry 

Camp, SE of Millaa Millaa, Hale 831968 (US). 

Myriotrema frustillatum Mangold spec. nov. ined. 

Type: Australia, New South Wales, Mt. Warning NP., track from summit to parking lot, Mangold 19 f 

(CANB-holotype; NSW-isotype). 

ETYMOLOGY – The epithet refers to the characteristic form of the isidia (from lat. 

frustillum =a small piece, a scrap). 


Fig. 70. Myriotrema frustillatum: growth habit (A), isidia (B), isidia and ascomata (C, D) and 

ascospore (E). A., C..: Hale 831415; B., E.: CANB-holotype; D.: Hale 831681. Bar= A: 2 mm; B: 

1 mm; C: 0.7 mm; D: 0.45 mm; E: 18 µm.



(pale)greenish-gray to pale yellowish-gray. Surface dull to slightly shiny, smooth, ±rugose to 

verrucose, distinctly to strongly fissured. Thallus covered by an incontinuous to continuous 

protocortex, up to c. 35 µm thick. Algal layer moderately to well developed, ±continuous, 

calcium oxalate crystals abundant, small to large, usually clustered. Isidia present, very 

characteristic, concolorous with thallus, irregularly shaped, up to c. 300 µm in diameter, with 

a depressed to stalk-like base, usually developing out of a pit-like structure (see fig. 70, B-D). 

Ascomata inconspicuous, usually very sparse or absent, moderately large, up to c. 800 µm in 

diam., roundish, perithecioid, solitary, becoming distinctly emergent, (irregular-) 

hemispherical to irregular-urceolate in older ascomata. Discs not visible from surface. Pores 

tiny, up to c. 70 µm in diam., roundish, entire, proper exciple not visible from surface. 

Thalline rim margin moderately thick, concolorous with thallus, thalline rim incurved. Proper 

exciple fused, moderately thick, pale yellowish to pale orange internally, orange to yellowishbrown 

or brownish marginally, weakly to distinctly amyloid at the base. Hymenium up to c. 

250 µm high, non-inspersed, strongly conglutinated, paraphyses thin, parallel, unbranched, 

straight to slightly bent in basal parts, distinctly curly in apical parts, tips slightly thickened, 

lateral paraphyses and columellar structures absent. Epihymenium indistinct, hyaline, without 

granules. Asci 1-2-spored, tholus and lateral walls evenly moderately thick in younger stages, 

thin when mature. Ascospores moderately to very large, densely eumuriform, cell walls and 

endospore thin, non-halonate, hyaline, non-amyloid, oblong to roundish-fusiform with 

roundish to narrowed-roundish ends, loci moderately small, ±angular, variably shaped, 

transverse septae thin, becoming somewhat irregular and indistinct with age, 80-200 x 20-38 

µm with multiple loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ orange-red, C-, Pd-; containing norstictic (major), connorstictic, 

subnorstictic and hyposalazinic (traces) acids. 


frustillatum grows on tree bark in (sub)tropical 


It is rare but wide-spread, occurring in northern 

Queensland and the Queensland/New South Wales 

border region and is currently only known from 

there. 

NOTES – Although it is predominantly sterile or 

is found with only very few ascomata, the new 

species is readily characterized by the presence of 

isidia and the presence of the norstictic acid 

chemosydrome. Fertile specimen are further 

characterized by perithecioid ascomata and large, 

densely eumuriform, hyaline, non-amyloid ascospores 

without thickened parts. Except for 

T. porinaceum, which has lateral paraphyses and 

lacks isidia, there are no similar species in 

Australia. The generic position of this taxon is 

uncertain. The brown outer proper exciple is 


M. frustillatum. 

reminiscent of Ocellularia, however, substrate structures are always absent. For the time 

being the new species is placed in Myriotrema until future phylogenetic studies become 

available.


SPECIMENS EXAMINED – Australia, Queensland: Atherton Tablelands: Lake Eacham NP., Mangold 29 j (F); 

Tumoulin Rd., 5 km from turnoff to Ravenshoe, Mangold 30 y pr.p. (F). Mapleton Falls, c.13 km W of 

Nambour, Hale 832002 (US). New South Wales: Tweed Range, Mebbin NP., 25 km SW of Murwillumbah, 

Mangold 21 p (F). Nightcap NP., Mnt. Nardi/Mnt. Matheson Track, Mangold 22 m, u (F). NW part of 

Wiangaree Forest Drive, N of Kyogle, Hale 831415 (US). Cambridge Plateau Forest Drive, 1-3 km N of picnic 

area, Richmond Range SF., 30 km W of Casino, Hale 831681 (US). 

Myriotrema glaucophaenum (Kremp.) Hale 

Mycotaxon 11: 133 (1980). Bas.: Thelotrema glaucophaenum Kremp., Nuov. Giorn. Bot. Ital. 7: 19 (1875). 

Ocellularia glaucophaena (Kremp.) Zahlbr., Cat. Lich. Univ. II: 591 (1923). Type: Borneo, 1866, Beccarii 92 

(M!-holotype, G-isotype). 

Ocellularia costaricensis Müll. Arg., Bull. Soc. Bot. Belgique 30: 75 (1891d). Rhabdodiscus costaricensis 

(Müll. Arg.) Vainio, Ann. Acad. Sci. Fenn. A 15(6): 184 (1921). Ocellularia alba f. costaricensis (Müll. Arg.) 

Redgr., Arkiv för Bot. 28A, 8: 18 (1936). Myriotrema costaricense (Müll. Arg.) Hale, Mycotaxon 11: 133 

(1980). Type: Costa Rica, Baie de Salinas, 1892, Pittier s.n. (Plantae costaricenses exsiccatae no. 5320) (Gholotype, 

M!-isotype). 

Thelotrema emergens Vain., Ann. Acad. Sci. Fenn., ser. A 15(6): 192 (1921a). Ocellularia emergens (Vain.) 

Zahlbr., Cat. Lich. Univ. II: 590 (1923). Type: Philippines, Mindanao, Butuan, C. M. Weber 1416 (TUR-Vain. 

26885!-holotype, NY!-isotype). 

Myriotrema subcostaricense Sipman, Acta Bot. Fenn. 150: 168 (1994). Type: Guyana, Upper Mazaruni 

District, Sipman & Aptroot 19424 (B-holotype, BRG-isotype). 


Fig. 72. Myriotrema glaucophaenum: growth habit (A), ascomata (B, C), ascospores (D, E) and 

ascospores showing amyloid reaction (F). A.: M-isotype of O. costaricensis; B.: Hale 831201; C.: 

Hale 831434; D., F.: Hale 832616; E.: TUR-holotype of T. emergens. Bar= A: 2 mm; B: 1 mm; C: 

0.75 mm; D: 9 µm; E: 7 µm; F: 8 µm.


Thallus epi- to hypophloedal, (moderately) thin, up to c. 200 µm high, pale yellowishgreen 

to (pale) olive. Surface ±shiny, smooth, continuous to rugose to verrucose, unfissured to 

slightly fissured. True cortex usually present, ±continuous, up to c. 25 µm thick, consisting of 

periclinal hyphae. Algal layer well to poorly developed, ±continuous, calcium oxalate crystals 

sparse to abundant, large and clustered, usually a distinct medulla layer present. Vegetative 

propagules not seen. Ascomata often conspicuous, becoming moderately large, up to c. 800 

µm in diam., in fused ascomata often larger, roundish to irregular, in particular in fused 

ascoma, apothecioid to somewhat chroodiscoid, solitary to more often and fused, becoming 

slightly to distinctly emergent, predominantly (irregular-) urceolate. Disc usually becoming 

partly to rarely entirely visible from surface, pale flesh colored, pruinose. Pores wide to 

gaping, up to c. 600 µm in diam., roundish to irregular, entire to slightly ragged, proper 

exciple often becoming apically to more rarely entirely visible from surface, ±free, whitish, 

sometimes slightly shrunken, incurved to erect to rarely slightly recurved. Thalline rim 

margin wide to gaping, rarely roundish to more often irregular, (moderately) thick, rarely 

entire to usually split to lacerate or eroded, rarely concolorous to distinctly whitish and 

somewhat pruinose due to exposed medulla, thalline rim usually becoming erect to slightly 

recurved. Proper exciple fused to partly or entirely free, thick, hyaline internally, pale 

brownish or grayish marginally, sometimes with inclusions of substrate particles, nonamyloid. 

Hymenium up to c. 100 µm high, non-inspersed, strongly conglutinated, paraphyses 

thick, irregular and often dist. septate, ±interwoven, with thickened, irregular tips, lateral 

paraphyses and true columella absent, columella-like structures often present due to fused 

ascomata. Epihymenium (moderately) thick, hyaline, with coarse grayish granules and fine 

crystals. Asci 8-spored, tholus moderately thick, thin when mature. Ascospores (very) small, 

transversely septate, sometimes with a single longitudinal septum, cell walls moderately thin 

to moderately thick, endospore (moderately) thick, covered by a thin halo, hyaline, variably 

amyloid, non-amyloid in younger ascospores, usually distinctly amyloid at maturity, 

±ellipsoid to oblong, with narrowed-roundish to subacute ends, loci ±roundish, subglobose to 

lentiform or oblong, end cells hemispherical to conical, septae moderately thin to moderately 

thick, often irregular, 10-18(20) x 5-8 µm with 2-6(8) (x2) loci. Pycnidia not seen, fide Frisch 

(2006) in thallus warts with brownish pore area, conidia fusiform, up to 6 x 1.5 µm. 

CHEMISTRY – Thallus K+ yellowish, C-, PD+ 

yellow; containing psoromic (major), 2'-0demethylpsoromic 

(minor to trace) and subpsoromic 

(trace) acids. 


glaucophaenum was collected in Australia on tree 

bark predominantly in tropical rainforests in 

altitudes ranging from 50 - 1200 m. It is a common 

species in Queensland. This is the first report for 

Australia, it is also known from the Neotropics 

(Frisch, 2006), Africa (ibid.), Sri Lanka (Hale, 

1981), Malaysia (ibid.) and the Philippines (ibid.) 


NOTES – This taxon is characterized by a ±shiny, 

corticate thallus, emergent, wide-pored and fused 

ascomata irregular paraphyses and small, 

transversely septate, hyaline ascospores and the 

presence of the psoromic acid chemosyndrome. 


M. glaucophaenum.


Similar Australian species include M. viridialbum, M. clandestinum and M. microporum, for 

differences see under these species. Myriotrema glaucophaenum is unusual since its ascomata 

are Thelotrema-like but it lacks lateral paraphyses. Frisch (2006) included M. glaucophaenum 

in a sub-group of Myriotrema, the M. viridialbum-group, which is, amongst other characters, 

outlined by the presence of a "massive, ±reticulate columella" (ibid.). Although 

morphologically as well as microscopically in the ascoma sections, columella-like structures 

are frequently present, it is obvious that these structures are not derived from single apothecia 

but formed of excipular remnants of merged ascomata. Thus, often two or more ascomata can 

be found in various stages of fusion in a single, or in two (three) fused (irregular) thalline 

rim(s). A feature that is, for example, also found in M. album (see also under this species). 

SPECIMENS EXAMINED – Australia, Queensland: Mt. Windsor logging area, just E of old Forestry Camp, NW 

of Mossman, Hale 831872, 832440, 832616 (US). Mt. Windsor, 5 km W of new Forestry Camp, NW of 

Mossman, Hale 830709 (US). Mt.Lewis Rd. 16 km N from Kennedy Hwy., W of Mossman, Hale 832596 (US). 

Atherton Tableland: Lake Euramoo, Lumbsch & Mangold 19127 e (F); Danbulla Forest Drive, E of Tinaroo: 

Lake Euramoo State Forest Park, Hale 831347 (US); Forest walk at Mobo Creek Crater, Hale 831434 (US); 1 

km W of Cathedral Fig Tree nr. Yungaburra, Hale 831201 (US); Along west boundary of Lake Eacham NP., on 

rd. from Atherton, Hale 831917 (US); Lake Eacham NP., track around lake, Hale 830284 (US); 22 km NE of 

Atherton, Mt. Haig, Tibell 15315 (UPS); Wongabel SF., S of Atherton on the Kennedy Hwy., Hale 831751 

(US); 13 km S of Ravenshoe on Tully Falls Rd., Hale 832631 (US). Blencoe Creek, Cardwell Range, 48 km NW 

of Cardwell, Streimann 36817 (B, CANB, US). About 7.5 km E of Wallaman Falls, W of Ingham, Hale 831966 

(US). Conway SF., 18 km E of Proserpine, Elix 20770 (CANB). Noosa NP., Palm Grove Track, Hale 830764 

(US).Solomon Islands, Santa Isabal Isl., Tatamba, Hill 11177 (US). French Guiana, Saül, Sipman 31637 (as 

exsiccate of M. subcostaricense - Lichenotheca Latinoamericana no. 134) (US). Costa Rica, Pitier 5321 (US). 

Myriotrema microporum (Mont.) Hale 

Mycotaxon 11: 134 (1980). Bas.: Thelotrema microporum Mont., Ann. Sci. Nat., Bot., sér. 3, 10: 130 (1848). 

Coscinedia micropora (Mont.) Massal., Atti I. R. Istit. Veneto, ser. 3, 5: 256 (1860). Ocellularia micropora 

(Mont.) Müll. Arg., Flora 74: 112 (1891e). Type: Indonesia, Java, Junghuhn s.n. (= Lichenes Javanicum no. 143) 

(PC-lectotype, selected by Hale [1981: 286]; FH-Tuck.!-, G-, H-Nyl.-, L-, M-, US!-isolectotypes). 

Thelotrema crassulum Nyl., Ann. Sci. Nat., Bot., sér. 4, 11: 258 (1859). Ocellularia crassula (Nyl.) Zahlbr., 

Cat. Lich. Univ. II(4): 588 (1923). Type: Mauritius ('Bourbon'), Boivin s.n. (PC-lectotype, selected by Hale 

[1981: 286]; M-, G-, H-Nyl. 22652-, H-Nyl.22653-isolectotypes). 

Ocellularia galactina Zahlbr., Ann. Crypt. Exot. 5: 216 (1932). Type: South Africa, Knysna, v.d. Byl 673 

(W-holotype; LD-, US!-isotypes). 


Thallus predominantly epi- to somewhat hypophloedal, thick, up to c. 800 µm high, color 

variable, predominantly (pale) olive in younger thalli becoming grayish or greenish-gray to 

yellowish-brown with age, often with a (dark-) gray hue. Surface dull to shiny, smooth, 

continuous to rugose, becoming distinctly fissured to areolate in mature stages. Cortex 

structures variable, often and particularly in younger thalli true cortex present, ±continuous, 

hyaline, up to c. 50 µm thick, consisting of periclinal to somewhat irregular hyphae, 

sometimes lacking a true cortex, then covered by a thin and often incontinuous protocortex. 

Algal layer well developed, continuous, calcium oxalate crystals lacking or sparse, then small 

and scattered to more rarely clustered, distinct medullar layer often present. Vegetative 

propagules not seen. Ascomata abundant, small, up to c. 250 µm in diam., roundish, 

apothecioid, solitary to marginally fused, predominantly immersed. Disc usually not visible 

from surface, very rarely becoming partly visible, pale flesh colored, epruinose. Pores small 

to rarely mod small, up to c. 150 µm in diam., roundish to irregular, predominantly ±split, 

proper exciple usually becoming apically to rarely entirely visible from surface, off-white, 

usually shrunken, incurved. Thalline rim margin becoming moderately wide to wide, roundish


Fig. 74. Myriotrema microporum: growth habit (A), ascomata (B), ascomata section (C), ascospores 

(D, E) and conidia (F). A.-D.: FH-isolectotype; E., F.: Lumbsch & Mangold 19174 a. Bar= A: 2.5 

mm; B: 1 mm; C: 125 µm; D: 7 µm; E: 2.5 µm; F: 3 µm. 

to somewhat irregular, (moderately) thin, entire to more often ±split, often ±distinctly raised, 

whitish or brighter than thallus, thalline rim incurved. Proper exciple usually becoming free in 

upper parts, (moderately) thin, with thin or lacking hyaline to pale yellowish area internally, 

entirely or marginally grayish to pale grayish-brown, sometimes faintly amyloid in lower 

parts. Hymenium up to c. 100 µm high, non-inspersed, moderately conglutinated, paraphyses 

somewhat irregular, ±interwoven, with slightly thickened tips, lateral paraphyses lacking, 

apical exciple sometimes forming lateral paraphyses-like structures due to radiating hyphae, 

columellar structures absent. Epihymenium indistinct to thin, hyaline, sometimes with fine 

grayish granules. Asci 8-spored, tholus moderately thick, thinning or not visible at maturity. 

Ascospores (very) small, transversely septate, cell walls and endospore (moderately) thick, 

non-halonate, hyaline, distinctly to strongly amyloid, oblong to ellipsoid to somewhat 

fusiform with roundish to subacute ends, loci roundish, oblong to lentiform, with same shaped 

to hemispherical to rarely conical end cells, septae moderately thin, regular, 10-16(18) x 5-8


µm with 3-4(5) loci. Pycnidia present, immersed, with inconspicuous dark pore, conidia 

fusiform, up to c. 5 x 2 µm. 

CHEMISTRY – Thallus K+ yellowish, C-, PD+ yellow; containing psoromic (major), 2'-0demethylpsoromic 

(minor to trace) and subpsoromic (trace) acids. 


microporum occurs on tree bark in (sub)tropical 

rainforests, rarely in wet sclerophyll forests in 

altitudes ranging from sea level to 1200 m. It is 

common and wide spread in the north-western 

Northern Territory and Queensland. This 

paleotropical species was also recorded from 

Africa, Mauritius, India (Nagarkar & al., 1988), Sri 

Lanka (Hale, 1981), SE Asia (eg., Hale, 1981; 

Nagarkar & al., 1988; Sipman, 1993), Samoa 

(Zahlbruckner, 1907) and Solomon Islands (Hale, 

1981). Records from the Neotropics are erroneous 

(Hale, 1981). 

NOTES – Mature and well developed specimen 

of M. microporum can be recognized morphologically 

by the thick, areolate thallus that becomes 

somewhat dull and (dark-)gray, and immersed 

ascomata with free exciple. It is further 

characterized by small, transversely septate, 


M. microporum. 

hyaline, amyloid ascospores with distinctly thickened parts and the psoromic acid 

chemosyndrome. In younger and poorly developed collections however, M. microporum can 

be easily confused with M. clandestinum, which differs by ascomata with fused exciple, an 

often exposed disc and slightly lager ascospores (up to 25 µm long) that sometimes have a 

single longitudinal septum. Other similar species include the psoromic acid containing M. 

glaucophaenum and M. temperatum, and M. olivaceum with different chemistry (for further 

differences to M. microstoma see under these species). The extratropical M. temperatum can 

be distinguished by a distinctly verruculose thallus with abundant calcium oxalate crystals 

and larger ascomata that have a fused proper exciple and an entire, thicker thalline rim 

margin. Myriotrema glaucophaenum differs by a thicker, never distinctly fissured or areolate 

thallus, larger, usually distinctly emergent ascomata with a thick, often distinctly split to 

lacerate thalline rim margin. 

SPECIMENS EXAMINED – Australia, Northern Territory: Curtain Falls, Litchfield Park, 38 km WSW of 

Batchelor, Elix 27577 (CANB). Kakadu NP., "Gungarre Monsoon Forest", A. & K. Kalb 30597 (hb Kalb). 

Channell Point, 23 km NNW of Daly River, Elix 27713 (CANB). Queensland: Uncertain locality, 1899, Bailey 

s.n. (BM-761745, -761745). Cape Tribulation Area, near the "Cape Tribulation Store", Mangold 33 v (F). Km 45 

on Mt. Windsor Rd., NW of Mossman, Hale 831418, 832300 (US). Mt. Windsor Tableland, 45 km NW of 

Mossman, Elix 16457 (B, CANB). Mt. Windsor logging area, just E of old Forestry Camp, NW of Mossman, 

Hale 831256, 831854, 832262 (US). Mt. Windsor, 5 km W of new Forestry Camp, NW of Mossman, Hale 

832719 (US). Mt. Windsor logging area, 9 km from rd. to old Forestry Camp and the main rd., Hale 830611 

(US). Mt.Lewis Rd., 10 km N from Kennedy Hwy., W of Mossman, Hale 831245, 831947, 832431 (US). 

Atherton Tablelands: Davies Creek Rd. 17 km S of Kennedy Hwy., S of Davies Creek Falls NP., E of Mareeba, 

Elix 85 (CANB, US), Hale 64024 (COLO, H, MEL) 831871, 832445, 832578, 832757 (US); Danbullah Drive, E 

of Tinaroo, forest walk at Mobo Creek Crater, Hale 831489, 832510 (US); 22 km NE of Atherton, Mt. Haig, 

Tibell 15313 (UPS); Lake Eacham NP.: Mangold 29 am (F), A. & M. Aptroot 22633 (ABL); Plath Rd. logging 

head, 9 km W of Plath Rd., off Kennedy Hwy, Herberton range, S of Atherton, Hale 830274, 831861, 832192 

(US); Malanda Falls, Lumbsch & Mangold 19132 zc (F); 15 km E of Malanda, close to Bora ground, Tibell


15369 (UPS); Tumoulin Rd., 5 km from turnoff to Ravenshoe, Mangold 30 k (F); S of hwy., 13 km E of jct. 

Kennedy Hwy. and Palmerston Hwy., E of Ravenshoe, Hale 831259; Arthur Bailey Rd., 9 km SSE of 

Ravenshoe, Streimann 46154 (B, CANB); Walter Hill Range, 26 km SE of Ravenshoe, Elix 17090 (CANB); 

Ravenshoe SF., Culpha Ck. Catchment, Cardwell Range, 41 km SE of Ravenshoe, Elix 16085, 16086 (CANB); 

Culpa logging area, 13 km from Koombooloomba rd. turnoff, SE of Tully Falls, Hale 832305 (US). Francis 

Range, Woopen Creek Rd., 25 km in from Bruce Hwy, NW of Innisfail, Hale 832085 (US). Dawson logging 

area, State Forest Reserve 605, 24 km S of Koombooloomba turnoff, WSW of Tully, Hale 831216, 831280, 

832110 (US). Cardwell Range: Blencoe Creek, 48 km NW of Cardwell, Streimann 36870 (CANB, H, US); 

Yuccabine Creek, Kirrima Rd., 27 km WNW of Cardwell, Elix 15745 (CANB); Kirrima SF., 24 km WNW of 

Cardwell, Elix 15666, Streimann 28584 (CANB). North Wallaman Logging Area, 36 km WNW of Ingham, 

Streimann 28799 (B, CANB). About 7.5 km E of Wallaman Falls, W of Ingham, Hale 831009 (US). Mt. Spec 

SF.: Paluma Range, 6 km SW of Paluma, Elix 20255 (B, CANB); Paluma to Hidden Valley Rd., 40 km S of 

Ingham, Streimann 64262 (B, CANB). Mt. Spec NP., Ridge on the Loop, on the Paluma Rd., WNW of 

Townsville, Hale 832419 (US). Conway Range NP., near Shute Harbour-Airlie Beach, Hale 832698, 832721 

(US). Cape Hillsborough NP., NW of Mackay, Hale 831292 (US). Eungella NP.: Tibell 14731 (UPS); NP. side 

rd. nr. Peases Lookout, off Darymple rd., Hale 832468 (US); Walking trail from Mt. Darymple rd. to Mt. 

Darymple summit, Rambold 4606 (M); Palm Walk Circuit, Hale 831125 (US); Trail from Broken River station 

to rainforest, Lumbsch & Mangold 19108 l (F); Finch Hatton Gorge, Lumbsch & Mangold 19116 d (F). Frasier 

Coast, River Heads, Lumbsch & Mangold 19092 o (F). Carabeen Nature Refuge, 45 km E of Warwick, Lumbsch 

& Mangold 19174 a (F). 

Myriotrema myrioporum (Tuck.) Hale 

Mycotaxon 11: 134 (1980). Bas.: Thelotrema myrioporum Tuck., Proc. Amer. Acad. Arts Sci. . 5: 412 

(1862). Ocellularia myriopora (Tuck.) Müll. Arg., Rev. Mycol. 10: 114 (1888). Type: Cuba, Monte Verde, 

Wright 129 (BM!-lectotype, selected here 10 ; FH-Tuck.-, US!-isolectotypes). 


Thallus epi- to hypophloedal, moderately thick, up to c. 200 µm high, (pale) olive to pale 

yellowish-gray. Surface slightly shiny, smooth, continuous to slightly verruculose, unfissured. 

True cortex usually present, predominantly continuous, slightly yellowish, thickness variable, 

up to c. 40 µm thick, formed of periclinal to irregular hyphae, sometimes partly not 

conglutinated forming a protocortex. Algal layer well developed, continuous, presence of 

calcium oxalate crystals variable, sparse to abundant, predominantly large, clustered, 

sometimes a distinct medallar layer present. Vegetative propagules not seen. Ascomata 

inconspicuous but often very abundant, predominantly ±small, up to c. 400 µm in diam., 

roundish, apothecioid, solitary to marginally fused, predominantly immersed to rarely 

somewhat emergent, then hemispherical. Disc often becoming partly visible from surface, 

pale flesh colored, epruinose to slightly pruinose. Pores moderately small, sometimes 

becoming wide, up to c. 200 µm in diam., predominantly roundish to rarely somewhat 

irregular, entire, proper exciple not visible from surface. Thalline rim becoming thick to very 

thick, sometimes funnel-shaped, brighter than thallus, forming a narrow fawnish to off-white 

ring-like area, thalline rim incurved. Inner ascomata margin two-layered, with a fused, 

(moderately) thin proper exciple and a thicker layer of conglutinated paraphyses, not clearly 

separated from the hymenium, pale yellowish, usually ±amyloid, turning reddish, often with a 

distinct, thick layer of grayish granules towards the epihymenium. Hymenium up to c. 70 µm 

high, non-inspersed, strongly conglutinated, paraphyses somewhat irregular, unbranched, 

often distinctly septate, slightly interwoven, with slightly to distinctly thickened, irregular 

tips, lateral paraphyses land columellar structures absent. Epihymenium indistinct to thin, 

hyaline, sometimes with fine grayish granules. Asci 8-spored, tholus moderately thin to 

10 The material in BM assigned as Wright 129 consists of several specimen of M. myrioporum, the upper, middle 

piece is here selected as type. In the US sample, the upper, larger specimen resembles the according isolectotype 

material.


lacking. Ascospores uniform, very small, with a single septum to more rarely not septated, 

cell walls and septum thick, non-halonate, hyaline, moderately amyloid, ellipsoid to fusiform, 

with narrowed-roundish to subacute ends, loci subglobose to (roundish-)conical, 10-13 x 4-6 

µm with 1- 2 loci. Pycnidia not seen. 

Fig. 76. Myriotrema myrioporum: growth habit (A), ascomata section (B) and ascospores (C, D). A.: 

US-isolectotype; B.: Hale 830606; C., D.: Hale 830011. Bar= A: 1.75 mm; B: 200 µm; C: 6 µm; 

D: 3 µm. 

CHEMISTRY – Thallus K-, C-, PD-; no secondary 



myrioporum was collected in Australia on tree bark 

in tropical rainforests in 900 m and an unknown 

altitude. It is a very rare species in Australia, 

occurring in northern Queensland. This is the first 

report for Australia and the Paleotropics, it is 

otherwise known from the Neotropics, indicating a 

pantropical distribution. 

N OTES – The 2-celled ascospores are 

characteristic of this species as well as the anatomy 

of the lateral exciple and hymenium, which was 

also mentioned by Redinger (1936). This ascomatal 

part shows a reddish amyloid reaction and often 


M. myrioporum.


includes grayish granules. It is morphologically similar to M. album and M. clandestinum but 

can be readily distinguished by the unique ascoma characters. Myriotrema uniseptatum from 

Panama is another similar species that differs in a pruinose thallus surface and smaller (up to 

7 x 3 µm) up to 3(4)-locular ascospores and in containing psoromic and stictic acid as major 

secondary metabolites. 


Portland Rds., Hale 830011 (US). About 8.5 km E of Wallaman Falls, W of Ingham, Hale 830606 (US). 

Myriotrema olivaceum Fée 

Méthode lichénographique et genera. Paris: tab. 1, fig. 25 (1824); Essai sur les cryptogames des écorces 

exotiques officinales. Paris: 103 (1825 ['1824']). Ocellularia olivacea (Nyl.) Müll. Arg., Mém. Soc. Phys. Hist. 

Nat. Genève 29(8): 7 (1887c). Ocellularia olivacea (Nyl.) Overeem, in Overeem-de Haas C. & Overeem-de 

Haas, D., Bull. Jard. Bot., sér. 3, 4: 118 (1922) [superfl. comb.]. Type: South America, "On Bonplandia 

trifoliata", s.c. (G-Fée 250-lectotype, selected by Hale [1974b: 24]; M-, PC-Mont.-isolectotypes). 

Thelotrema subcrassulum Vain., Mycologia 21: 38 (1929). Ocellularia subcrassula (Vain.) Zahlbr., Cat. 

Lich. Univ. 8: 245 (1932). Type: Puerto Rico, near El Yunque, Fink 725 (MICH-lectotype, selected by Hale 

[1978: 12]; US!-isolectotype). 


Thallus predominantly epi- to somewhat hypophloedal, thick, up to c. 600 µm high, color 

variable, predominantly (pale) olive in younger thalli, becoming grayish or greenish-gray to 

yellowish-brown with age, sometimes with a (dark-) gray hue. Surface dull to shiny, smooth, 

continuous to rugose, becoming distinctly fissured to areolate. Thallus cover variable, often 

and particularly in younger thalli true cortex present, continuous to incontinuous, hyaline, up 

to c. 40 µm thick, consisting of periclinal to somewhat irregular hyphae, sometimes lacking a 

true cortex, then covered by a thin and often incontinuous protocortex. Algal layer well 

developed, continuous, calcium oxalate crystals lacking or sparse, then ±small and clustered, 

a distinct medullar layer present. Vegetative propagules not seen. Ascomata becoming 

abundant, small, up to c. 300 µm in diam., roundish, apothecioid, solitary to marginally fused, 

predominantly immersed to slightly raised. Disc not visible from surface to very rarely 

becoming partly visible, pale flesh colored, epruinose. Pores small to more rarely moderately 

small, up to c. 100 µm in diam., roundish to irregular, predominantly ±split, proper exciple 

usually becoming apically to rarely entirely visible from surface, off-white, usually shrunken, 

incurved. Thalline rim margin becoming moderately wide to wide, roundish to somewhat 

irregular, thin to moderately thick, entire to ±split, often ±distinctly raised, whitish or brighter 

than thallus, thalline rim incurved. Proper exciple usually becoming free in upper parts, 

(moderately) thin, with thin hyaline to pale yellowish area internally, orange to pale brown 

marginally, in older stages often (dark) brown apically, sometimes faintly amyloid in lower 

parts. Hymenium up to c. 90 µm high, non-inspersed, moderately conglutinated, paraphyses 

somewhat irregular, ±interwoven, with slightly thickened tips, lateral paraphyses absent, 

apical exciple sometimes forming lateral paraphyses-like structures due to radiating hyphae, 

columellar structures absent. Epihymenium indistinct, hyaline, without granules. Asci 8spored, 

tholus moderately thick, thinning or not visible at maturity. Ascospores very small, 

transversely septate, cell walls and endospore (moderately) thick, non-halonate, hyaline, 

distinctly to strongly amyloid, oblong to ellipsoid to somewhat fusiform in younger stages, 

with roundish to narrowed-roundish to subacute ends, loci roundish to slightly angular, 

subglobose to oblong to lentiform, with same shaped to more hemispherical or rarely conical 

end cells in younger stages, septae moderately thin, regular, 10-15 x 5-7 µm with 3-4(5) loci. 

Pycnidia not seen.


Fig. 78. Myriotrema olivaceum: growth habit (A, B), ascomata (C), ascomata and thallus section (D), 

ascospores showing amyloid reaction (E) and ascospore (F). A.: Hale 832174; B,. D.-F.: 

Streimann 30299; C.: US-isolectotype of T. subcrassulum. Bar= A, B: 1 mm; C: 0.75 mm; D: 125 

µm; E: 7 µm; F: 2 µm. 

CHEMISTRY – Strain I: Thallus K+ red, C-, PD-; containing olivaceic (major) and norsubnotatic 

(major to minor) acid. Strain II: Thallus K(+) reddish, C-, PD-; containing O-methylolivaceic 

and norisonotatic (majors) and olivaceic (minor) acid. 

ECOLOGY AND DISTRIBUTION – Myriotrema olivaceum grows on tree bark in (sub)tropical 

rainforests and more often in moist habitats as coastal forests, mangroves or monsoon forests 

in predominantly lower altitudes ranging from sea level to 400 m. Four specimens were 

collected at 1200 m. It is a common and wide-spread species occurring in north-western 

Northern Territory and Queensland. This is the first report for Australia. Previously it was 

recorded from the Neotropics, Africa (Frisch, 2006), Sri Lanka (Hale, 1981), Andaman


Islands (Nagarkar & al., 1986), New Caledonia 

(Hale, 1981) and Marquesas Islands (ibid.) 

indicating a pan(sub)tropical distribution. 

NOTES – This species is morphologically and 

anatomically almost indistinguishable from 

M. microporum. However, some minor differences 

in thallus thickness, ascomata characters and 

ascospore size (in M. microporum the thallus is up 

to 800 µm thick, the exciple grayish and less 

pigmented, ascospores up to 18 x 8 µm) were found 

in the present material. Further, the two species 

differ chemically (psoromic acid chemosydrome in 

M. microporum). The relationships of the two 

species require additional studies, including 

molecular data. 

Unfortunately no type material of M. olivaceum 

was available for study. Three examined specimen, 

all from the same location (Mt. Lewis area, SW of 

Mossman - Elix 16919, 16928, Streimann 30299) 


M. olivaceum. 

are assigned to the chemical strain II. They chemically agree with Frisch's (2006) description 

of M. subterebrans from Africa. The type collection was also not available for study, hence 

no further taxonomic statement can be made. Besides olivaceic acid in lower concentrations, 

strain II contains O-methylolivaceic and norisonotatic acid, with identical Rf-values and spotcharacters 

as described by Frisch (ibid.) for the unknown compounds 'neoterebrans unknown' 

and 'olivaceum unknown high' reported to occur as major compounds in M. neoterebrans. 

Both latter mentioned compounds are chemically closely related to olivaceic acid. Since strain 

II is morphologically identical to chemotype I, it is treated as a chemotype until the type 

material of M. subterebrans as well as M. olivaceum becomes available. 

Thelotrema configuratum 11 from Sarawak (Malaysia) is a similar species, which contains 

only norsubnotatic acid, has small, immersed ascomata with a free exciple, and similar 

ascospores. It was combined to Myriotrema (Hale, 1980) and thus, could be expected to be 

closely related to M. olivaceum (its chemical relationship was mentioned by Hale, 1974). 

However, the examination of the type collection revealed conspicuous lateral paraphyses (up 

to 20 µm long), indicating that the species belongs to Thelotrema. 

SPECIMENS EXAMINED – Strain I: Australia, Northern Territory, Litchfield NP., 100 km S of Darwin, A. & K. 

Kalb 34256 (hb. Kalb). Queensland: Cape Tribulation Area: Cape Tribulation Beach, Hale 830306 (US); C. 40 

km N of Mossman, A. & K. Kalb 27050 (hb. Kalb). Mossman Gorge NP., near Mossman, Hale 830637 (US). 

Main Coast Range, 19 km NNW of Mt. Molloy, Streimann 30299 (B). Crystal Cascades, 5 km W of Cairns, 

Lumbsch & Mangold 19120 f (F). Boat Ramp c. 2 km E of Edmonton, SE of Cairns, Lumbsch & Guderley 

11139 d (F). Little Crystal Creek Falls, Mt.Spec NP., the falls on Paluma Rd., Hale 831218 (US). Murray Falls, 

W of Kennedy, Hale 832547 (US). Conway NP., 22 km NE of Proserpine: At the campground 0.5 km E of 

Shute Harbor, Tibell 14663, 14671 (UPS); Near Shute Harbour-Airlie Beach, Hale 832174 (US). Conway 

Peninsula, E of Proserpine, Scott 622 (BRI). Eungella NP., Finch Hatton Gorge, Lumbsch & Mangold 19113 f 

(F). Frasier Coast, River Heads, Lumbsch & Mangold 19092 g (F). Amama SF., 6 km SW of Amamoor, S of 

Glympie, Hale 831365 (US). 

Strain II: Australia, Queensland: Main Coast Range, track to Mt. Lewis, 19 km NNW of Mt. Molloy, Elix 

16919, 16928, Streimann 30299 (CANB). 

11 Thelotrema configuratum (Hale) Mangold comb. nov. ined. Bas.: Ocellularia configurata Hale, Phytologia 

27: 491 (1974). Myriotrema configuratum (Hale) Hale, Mycotaxon 11: 133 (1980). Type: Malaysia, Borneo, 

Sarawak, N of Sibu, Hale 31207 (US!-holotype).


Myriotrema phaeosporum (Nyl.) Hale 

Mycotaxon 11: 134 (1980). Bas.: Thelotrema phaeosporum Nyl., Ann. Sci. Nat. Bot., sér. 4: 242 (1859). 

Leptotrema phaeosporum (Nyl.) Muell.Arg., Flora 65: 499 (1882). Type: Papua New Guinea, New Ireland, Port 

Carteret, Lesson s.n. (G!-, US!-isotypes). 

Leptotrema polyporum Riddle, Mycologia 15: 79 (1923). Type: Isle of Pines (New Caledonia), Sierra de los 

Caballos, Jennings 229 a (FH-lectotype, selected by Hale [1972 in herb.], NY!-isolectotype). 


Fig. 80. Myriotrema phaeosporum: growth habit (A), ascomata (B-D), ascoma and thallus section (E) 

and ascospores (F, G). A., B.: Bailey 70; C., D.: Elix 20878; E.-G.: G-isotype. Bar= A: 2 mm; B: 1 

mm; C: 1.25 mm; D: 0.3 mm; E: 200 µm; F, G: 6 µm.


Thallus predominantly epiphloedal, (moderately) thick, up to c. 1 mm high, bulging and 

flaking away from the substrate, pale greenish gray to pale yellowish-green. Surface ±dull, 

smooth, continuous, with a grainy-speckled pattern, unfissured to sometimes slightly fissured. 

True cortex absent, thallus covered by a continuous to incontinuous protocortex up to c. 30 

µm thick. Algal layer usually well developed, continuous, sometimes becoming incontinuous 

due to crystal inclusions, calcium oxalate crystals abundant, predominantly large, scattered or 

more often clustered, sometimes arranged in columns. Vegetative propagules not seen. 

Ascomata inconspicuous, (moderately) small, up to c. 400 µm in diam., roundish, 

predominantly perithecioid, solitary, immersed. Disc not visible from surface. Pores tiny to 

small, up to c. 100 µm in diam., roundish, entire, apical proper exciple becoming visible from 

the outside, forming a fused to indistinctly free inner pore margin, incurved, brighttranslucent 

to pale brownish, in older ascomata often thinned, then appearing more dark. 

Thalline rim margin moderately thick, roundish, entire, often brighter then thallus, thalline 

rim not distinguishable from surface. Proper exciple fused to apically slightly detached, 

moderately thin to moderately thick, pale yellowish internally, pale orange marginally, nonamyloid. 

Hymenium up to c. 150 µm high, non-inspersed, weakly conglutinated, paraphyses 

±bent to distinctly curly in apical parts, sometimes with ±distinct septation, moderately 

interwoven, with unthickened to slightly thickened tips, lateral paraphyses and columellar 

structures absent. Epihymenium indistinct or lacking, hyaline, without granules. Asci 8spored, 

tholus distinctly narrowed, thin to not visible at maturity. Ascospores (very) small, 

(sub-)muriform, cell walls thick, endospore moderately thin, non-halonate, becoming 

distinctly brown at early maturity, faintly to distinctly amyloid only in younger ascospores, 

oblong to predominantly ellipsoid to rarely somewhat fusiform, with rounded to narrowedrounded 

to rarely subacute ends, loci roundish to slightly angular, oblong to subglobose or 

somewhat irregular, with same shaped end cells, transverse septae moderately thin to 

moderately thick, regular to irregular, 10-25 x 7-15 µm with 4-8 x 1-4(-5) loci. Pycnidia not 

seen. 

CHEMISTRY – Thallus K+ yellow, C-, PD+ orange; containing constictic and stictic acids 



phaeosporum was collected in Australia on tree 

bark in (sub)tropical rainforests, wet sclerophyll 

forests and mangroves in altitudes ranging from sea 

level to 600 m. It is a moderately common and 

wide-spread species, occurring in northern 

Queensland, Norfolk Island and Lord Howe Island 

(New South Wales). This paleotropical species was 

reported from the Philippines (Vainio, 1921), 

Papua New Guinea and New Caledonia. 

NOTES – This taxon is characterized by a ±dull, 

smooth, bulging and thick thallus that lacks red 

crystals, immersed, perithecioid, small-pored 

ascomata with a predominantly fused exciple, 

brown, muriform small ascospores and the presence 

of the stictic acid chemosyndrome. It is similar to 

L. wightii, which can be readily distinguished by 

the presence of red anthraquinone crystals in the 

thallus, larger, more open ascomata, more distinctly 

Fig. 81 . Australian distribution of 

M. phaeosporum.


globose, less oblong ascospores with fewer loci (up to 6 x 4) that are irregularly arranged, and 

the absence of the stictic acid chemosyndrome. Also similar is L. subcompunctum, see under 

that species for differences. Other similar, brown-spored, stictic acid containing taxa include 

L. compunctellum, M. desquamans and M. trypaneoides. They can be readily distinguished by 

larger ascospores (up to 130 µm in L. compunctellum, up to 35 µm in M. desquamans, up to 

40 µm in M. trypaneoides), the presence of a true cortex in the latter two taxa, and an 

inspersed hymenium in M. trypaneoides. 

SPECIMENS EXAMINED – Australia, Queensland: Iron Range, Hill 1035497 (MEL). Thornton Range, NW of 

Mossman, Hale 832212 (US). Cairns, 5 km N of the city near the airport, A. & M. Aptroot 22173 (ABL). Clark 

Range, 46 km SSW of Proserpine, Elix 20878 (CANB). Wilson Beach, 17 km SE of Proserpine, Elix 20980 

(CANB). Whitsunday Islands, Long Island, N of Happy Bay, Tibell 14643 (UPS). Conway Range NP., near 

Shute Harbour-Airlie Beach, Hale 830972 (US). Noosa NP., E of Noosa Heads, Hafellner 16729 (GZU). 

Kondalilla Falls, SW of Nambour, Hale 832551 (US). Moreton Bay, Mt.Cooluma, Windolf 694631 (BRI). 

Blackall Ranges, Wilson s.n. (NSW-539380). Upper Coomera, Wilson s.n. (NSW-539384). Brisbane, Bailey 70 

(BM). Norfolk Island: Mt. Pitt Reserve, Red Road Track to Mt. Bates, Elix & Streimann 18614 (B), Elix 18855 

(CANB). New South Wales, Lord Howe Island: Max Nicholls Track, Elix 32717 (CANB); Jct. of Kims Lookout 

and Max Nicholls Tracks, Elix 42061 (CANB); Track from Smoking Tree Ridge to Rocky Fun, Elix 42461 

(CANB); Goat House Cave, Elix 42096, 42211 (CANB); Trail to Goat House Cave, Weber & Colson L-72185 

(COLO, US). Uncertain locality: 'Bailey Book', p. 21, s.c. (BRI-AQ720151); 'Shirley Book', p. 22, s.c. (BRI- 

AQ721239). 

Myriotrema polytretum Hale 

Bull. Br. Mus. nat. Hist. (Bot.) 8(3): 291 (1981). Type: Sri Lanka, Sabaragamuwa, Kegalla, Hale 50238 

(US!-holotype, BM-isotype). 



(pale) olive to yellowish-gray to grayish, sometimes with pale patches due to a partly absent 

algal layer. Surfaces shiny to velvety, smooth, continuous to slightly rugose, unfissured. True 

cortex usually present, predominantly continuous, slightly yellowish, thickness variable, up to 

c. 30 µm thick, consisting of periclinal to irregular hyphae. Algal layer moderately well 

developed, continuous but sometimes partly thinning or lacking (see above), calcium oxalate 

crystals moderately abundant, small to large, clustered to more rarely scattered, sometimes a 

distinct medullar layer present. Vegetative propagules not seen. Ascomata usually 

inconspicuous, ±small, up to c. 350 µm in diam., roundish, apothecioid, solitary to marginally 

to rarely entirely fused, often clustered, forming patches of densely arranged ascomata, 

immersed. Disc often becoming partly visible from surface, pale flesh colored, epruinose to 

slightly pruinose. Pores (moderately) small to sometimes becoming wide, up to c. 200 µm in 

diam., predominantly roundish to somewhat irregular, entire to rarely slightly split, proper 

exciple not visible from surface. Thalline rim margin (moderately) thin, brighter than thallus, 

forming a narrow to moderately wide fawnish to more often off-white ring-like area, level 

with thallus to often somewhat raised, thalline rim incurved. Proper exciple fused, 

(moderately) thin, hyaline to pale yellowish internally, pale orange to yellowish-brown 

marginally, sometimes grayish-brown apically, non-amyloid. Hymenium up to c. 70 µm high, 

non-inspersed, strongly conglutinated, paraphyses straight to slightly bent, ±irregular and 

distinctly septate, parallel to slightly interwoven, with distinctly thickened, somewhat 

irregular tips, lateral paraphyses and columellar structures absent. Epihymenium indistinct to 

thin, hyaline, sometimes with fine grayish granules. Asci 8-spored, tholus (moderately) thick, 

not visible at maturity. Ascospores (very) small, transversely septate, cell walls (moderately) 

thick, endospore moderately thick, sometimes with thin halo, hyaline, predominantly


distinctly to strongly amyloid, oblong to ellipsoid to somewhat fusiform, with roundish to 

narrowed-roundish ends, loci predominantly roundish, subglobose to oblong to slightly 

irregular, with same shaped or hemispherical to conical end cells, septae thin to moderately 

thick, ±regular, 10-18(22) x 6-8 µm with 3-4 loci. Pycnidia not seen. 

Fig. 82. Myriotrema polytretum: growth habit (A), ascomata (B), ascoma section (C), ascospores (D) 

and same ascospores showing amyloid reaction (E). A., B.: Mangold 29 n; C.-E.: US-holotype. 

Bar= A: 1 mm; B: 0.5 mm; C: 150 µm; D, E: 5 µm. 

CHEMISTRY – Thallus K-, C-, PD+ yellowish to reddish; containing 2-hydroxyhypoprotocetraric 

(major), convirensic, 2- hydroxynornotatic and hypoprotocetraric (traces) 

acids. 

ECOLOGY AND DISTRIBUTION – Myriotrema polytretum was collected in Australia on tree 

bark in tropical rainforests. It is extremely rare, being restricted to northern Queensland. This 

is the first report for Australia. Previously this paleotropical taxon was recorded from Sri 

Lanka, Malaya and Sarawak (Hale, 1981). 

NOTES – This taxon is characterized by its unusual thallus chemistry, otherwise it is very 

similar to M. clandestinum (with the psoromic acid chemosyndrome) and M. album (lacking 

secondary compounds). With these species it has a corticate, unfissured thallus in common 

with small, immersed ascomata with a fused proper exciple and small, transversely septate,


hyaline, amyloid, thick-walled ascospores. 

2-hydroxyhypoprotocetraric acid is usually found 

as an accessory compound of hypoprotocetraric 

acid, only one other species, the otherwise also 

quite similar O. thwaitesii 12 from Sri Lanka is 

known to contain 2-hydroxyhypoprotocetraric acid 

as a major compound, too. It differs, however, by a 

slightly carbonized exciple and ascospores that 

have up to 7, distinctly oblong loci (up to 22 x 7 

µm in size). Myriotrema microporellum, known 

from Africa and the Neotropics is also similar and 

contains hypoprotocetraric acid as major compound 

and 2-hydroxyhypoprotocetraric acid only in traces. 

However, it also differs in slightly larger 

ascospores (up to 24 µm, with up to 6 loci). Several 

other similar taxa with hypoprotocetraric acid as a 

major compound all differ from M. polytretum by a 

±distinctly free proper exciple: M. glauculum, M. 

plurifarium, M. congestum and M. neofrondosum. 


Portland Rds., Hale 832650 (US). Atherton Tablelands, Lake Eacham NP., Mangold 29 n (F). Malaysia, 

Sarawak, Sibu, Hale 30434 (US). Malaya, Selangor, Hale 30271 (US). 

Myriotrema protoalbum Hale 

Bull. Br. Mus. nat. Hist. (Bot.) 8(3): 292 (1981). Type: Sri Lanka, Western Province, Kalatura, Hale 51044 

(US!-holotype, BM-isotype). 



M. polytretum. 

Thallus epi- to hypophloedal, (moderately) thin, up to c. 100 µm high, (pale) grayishgreen. 

Surface shiny, smooth, continuous, unfissured. True cortex present, continuous to 

incontinuous, up to c. 30 µm thick, consisting of periclinal hyphae. Algal layer moderately 

well developed, continuous to incontinuous, presence of calcium oxalate crystals variable, 

abundant to lacking, small to large, clustered, sometimes forming short layers. Vegetative 

propagules not seen. Ascomata variable, inconspicuous, small to very small, up to c. 250 µm 

in diam., roundish to elongate to sometimes distinctly lirelliform (then up to 500 µm long), 

apothecioid, solitary to marginally fused, often arranged in rows, immersed to slightly raised, 

then hemispherical. Disc rarely becoming visible from surface, pale flesh colored, epruinose. 

Pores predominantly small to very small, up to c. 80 µm in diam. (wider in elongate 

ascomata), roundish to often irregular or elongate, entire to coarsely split, proper exciple not 

visible from surface. Thalline rim margin (moderately) thin, variable in color, often brighter 

than thallus to whitish, sometimes brownish, rarely concolorous with thallus, thalline rim 

incurved. Proper exciple fused, predominantly moderately thin, pale yellowish internally, 

yellowish-brown to brownish marginally, basally often amyloid, sometimes amyloidity 

extending throughout the subhymenium. Hymenium up to c. 90 µm high, non-inspersed, 

moderately conglutinated, paraphyses ±straight to irregular, slightly interwoven, with 

±distinctly thickened, somewhat irregular tips, lateral paraphyses and columellar structures 

12 Ocellularia thwaitesii (Hale) Mangold comb. nov. ined. Bas.: Myriotrema thwaitesii, Bull. Br. Mus. nat. Hist. 

(Bot.) 8(3): 295 (1981). Type: Sri Lanka, Sabaragamuwa, Kegalla, Hale 50221 (US!-holotype, BM-isotype).


Fig. 84. Myriotrema protoalbum: growth habit (A), ascomata (B, C), ascoma section (D), ascus (E), 

younger ascospore (F) and mature ascospore (G). A., C., E.-G.: Mangold 39 r; B., D.: USholotype. 

Bar= A: 1 mm; B: 0.5 mm; C: 0.6 µm; D: 100 µm; E: 5 µm; F: 4 µm; G: 3 µm. 

absent. Epihymenium indistinct, hyaline, without granules. Asci 8-spored, tholus 

(moderately) thick, sometimes with distinct, tapered ocular chamber, thin when mature. 

Ascospores (very) small, transversely septate, rarely with a weakly developed, single 

longitudinal septum, cell walls (moderately) thick, endospore moderately thin to moderately 

thick, variably halonate, especially in younger ascospores sometimes with distinct, thick halo, 

in older ascospores often indistinct or lacking, hyaline, non-amyloid to faintly amyloid, 

predominantly oblong to ellipsoid, with roundish to narrowed-roundish ends, loci roundish to 

more rarely somewhat acute, subglobose to oblong to rarely irregular, usually with same


shaped end cells, septae moderately thin to often (moderately) thick, regular to irregular, 10- 

22 x 5-9 µm with 3-4(5) (x2) loci. Pycnidia not seen. 



protoalbum was collected in Australia on tree bark 

in tropical rainforests in altitudes ranging from 60 

to 1500 m. It is a rare species in northern 

Queensland. This is the first report for Australia. 

Previously it was only known from Sri Lanka. 


corticate thallus, small, ±immersed ascomata that 

are arranged in rows and sometimes are distinctly 

elongate, the small, hyaline, transversely septate 

ascospores, and the absence of secondary 

compounds. Myriotrema album is a similar species 

that can be distinguished by a thicker thallus, 

somewhat larger, distinctly amyloid ascospores and 

ascomata that are never elongate. The Australian 

collections differ from the type in having a thicker 

thallus, a never brownish thalline rim margin and 

ascomata that can be often found slightly to 

distinctly elongate. The typical row-like 

arrangement of the ascomata, however, was already mentioned in the original species 

description (Hale, 1981). The tholus of M. protoalbum has an often distinct, tapered ocular 

chamber (Fig. 85, E). A similar tholus structure was reported for two Chapsa species, C. 

eitenii and C. zahlbruckneri (Frisch, 2006). 

SPECIMENS EXAMINED – Australia, Queensland: Daintree NP., Mossman Gorge Section, near western border 

of National Park along Mossman creek, Mangold 36 b (F). Wooroonooran NP., Bellenden Ker Range, centre 

peak, 7 km W of Bellenden Ker, A. & M. Aptroot 22503 (ABL). Atherton Tablelands, Lake Eacham NP., 

Mangold 29 ar (F). Babinda Boulders, Mangold 39 r (F). Eungella NP., near Pease's Lookout, Hale 831699 

(US). Sri Lanka, Sabaragamuwa: Ratnapura, Hale 51219 (US); Kegalla, Hale 50247 (US). 

Myriotrema rugiferum (Harm.) Hale 

Mycotaxon 11: 135 (1980). Bas.: Thelotrema rugiferum Harm., Bull. Séanc. Soc. Sci. Nancy, ser. 3, 13: 44 

(1912). Type: New Caledonia, Pionniero 42 (DUKE!-lectotype, selected by Hale [1981: 292]; S!-isolectotype). 

Thelotrema subcaesium Nyl., Flora 52: 120 (1869). Thelotrema concretum var. subcaesium (Nyl.) Redgr., 

Arkiv för Bot. 28A, 8: 96 (1936). Type: Brazil, Rio de Janeiro, Glaziou 2193 (H-Nyl.-lectotype, selected by 

Hale [1974a: 32]; C!-, US!-isolectotypes). 

Myriotrema multicavum Hale, Bull. Br. Mus. nat. Hist. (Bot.) 8(3): 288 (1981). Type: Sri Lanka, Southern 

Province, Galle Distr., Hale 46150 (US!-holotype). 



M. protoalbum. 

Thallus epi- to hypophloedal, predominantly moderately thick, up to c. 400 µm high, (pale) 

olive to (pale) grayish-green or yellowish-gray to grayish. Surface variable, velvety to shiny, 

smooth, continuous to more often rugose and/or distinctly verruculose, unfissured to slightly 

fissured or coarsely cracked. True cortex usually present, predominantly continuous, slightly


yellowish, thickness variable, up to c. 30 µm thick, consisting of periclinal hyphae, sometimes 

partly not conglutinated forming a protocortex. Algal layer well developed, continuous, 

presence of calcium oxalate crystals variable, sparse to very abundant, predominantly large, 

Fig. 86. Myriotrema rugiferum: growth habit (A, B), ascomata (C), pycnidia (D), ascomata section 

(E), pycnidium section (F), ascospores (G), ascospore showing amyloid reaction (H) and conidia 

(I). A., G.: DUKE-lectotype; B. C., E., H.: Lumbsch & Mangold 19116 a; D., F., I.: Lumbsch & 

Mangold 19116 h. Bar= A, B: 1 mm; C, D: 0.3 mm; E: 100 µm; F: 50 µm; G: 8 µm; H: 7 µm; I: 6 

µm.


clustered, distinct medullar layer present. Vegetative propagules not seen. Ascomata 

inconspicuous but often very abundant, small, up to c. 400 µm in diam., roundish to irregular 

in fused ascomata, apothecioid, solitary to fused, predominantly immersed to rarely somewhat 

emergent, then hemispherical to cone-shaped. Disc usually not visible from surface, 

sometimes becoming partly visible, pale flesh colored, distinctly to strongly pruinose. Pores 

(moderately) small, sometimes becoming wide or gaping, up to c. 100(200) µm in diam., 

predominantly irregular and ±split, proper exciple becoming apically to more rarely entirely 

visible from surface, whitish, shrunken, incurved. Thalline rim margin becoming moderately 

wide to wide or gaping, roundish to somewhat irregular, (moderately) thin, predominantly 

entire to slightly split, whitish, thalline rim incurved to very rarely somewhat erect. Proper 

exciple free in upper parts, thin, hyaline internally to brownish or grayish-brown marginally, 

often with an apical ±thick layer of grayish granules, non-amyloid. Hymenium up to c. 

90(110) µm high, non-inspersed, strongly conglutinated, paraphyses bent, unbranched to 

slightly branched towards the exciple, moderately to slightly interwoven, with slightly to 

distinctly thickened, irregular tips, lateral paraphyses and true columella absent, in fused 

ascomata columella-like structures sometimes present. Epihymenium predominantly 

(moderately) thick, hyaline, with fine to more often coarse grayish granules. Asci (4-)8spored, 

tholus thick, thin when mature. Ascospores small, submuriform, cell walls thick, 

endospore (moderately) thick, non-halonate, hyaline, distinctly to strongly amyloid, 

subglobose to oblong or ellipsoid to more rarely fusiform, with roundish to narrowedroundish 

to rarely subacute ends, loci roundish to slightly angular, subglobose to oblong or 

somewhat irregular, with hemispherical to same shaped end-cells, transverse septae distinct, 

moderately thin, regular to irregular, 15-25 x 7-10 µm with 4-6(7) x 1-3(4) loci. Pycnidia 

present, in thallus warts, often crowded, with colorless to dark, often wide pore surrounded by 

a bright area, conidia fusiform to somewhat irregular, up to c. 5 x 1.5 µm. 


(major to minor) and subpsoromic (trace) acid. 


rugiferum occurs on tree bark in (sub)tropical 

rainforests, coastal forests and mangroves, rarely 

also in wet sclerophyll forests in altitudes ranging 

from sea level to 1200 m. This common and widespread 

species occurs in Queensland and northern 

New South Wales. This is the first report for 

Australia. It is pantropical and has been recorded 

from the Neotropics, India (Hale, 1981), Sri Lanka, 

Philippines (Ibid.), New Caledonia and Solomon 

Islands (Ibid.). 

N OTES – Myriotrema rugiferum is 

morphologically variable, but all samples agree in 

having a thick, corticate thallus, small, predominantly 

immersed ascomata with a visible free 

exciple, small, hyaline, submuriform, thick-walled, 

amyloid ascospores and containing the psoromic 

acid chemosydrome. Similar is M. viridialbum; for 

differences see under this species. The morpho- 


M. rugiferum. 

logical extremes of M. rugiferum were previously regarded as different species, but are 

connected by intermediate forms. One of the morphological extremes resembles the type of


T. subcaesium, with a velvety, continuous and unfissured thallus, few calcium oxalate crystals 

and grouped, predominantly solitary ascomata with only indistinctly visible free proper 

exciple. The more common morph, in which agrees with the New Caledonian type, has a 

shiny, incontinuous, thallus with abundant, usually large and clustered crystals. 

A similar Australian species with small, submuriform, hyaline, amyloid ascospores that 

lacks thallus chemistry is M. subconforme, for further differences see under this species. A 

similar psoromic acid containing taxon known from South America and Borneo (Hale, 1978) 

is M. concretum. The type material of M. concretum was not available, but is distinguished 

from M. rugiferum by a strongly fissured, dull thallus and urceolate ascomata with fused 

exciple. 


Portland Rds., Hale 832747 (US). Browns Creek, Lockhart River Settlement rd., 30 km SW of Cape Weymouth 

(18 km NW of Lockhart River Settlement), Streimann 56457 (B, CANB). Cape Tribulation Area: Cape 

Tribulation Beach, Hale 831746 (US); Track to Cape Tribulation Beach, Mangold 32 ze (F); Myall Beach, 

Lumbsch & Mangold 19160 n, s, x, 19161 f, j, u, Mangold 31 f (F); Noah Head (Mairdja Botanical Walk), 

Streimann 45784 (CANB); Cow Bay, Streimann 45991 (B, CANB); 2 km W of main rd. between Oil Palms and 

Coopers Creek, Hale 832104 (US). Thornton Range, at tower turnoff on CREB rd (to Cooktown), c.15 km N of 

the Daintree River crossing, NW of Mossman, Hale 832144 (US). Near Daintree, along Daintree River, A. & K. 

Kalb 27051 (hb. Kalb). Mt. Windsor Tableland, 38-45 km NW of Mossman, Elix 16432, 16526, 16548 (CANB). 

Mt. Windsor, 5 km W of new Forestry Camp, NW of Mossman, Hale 831868, 832598, 832611, 832644 (US). 

Mt. Windsor logging area, near jct. rd. to old Forestry Camp and main rd., NW of Mossman, Hale 831859 (US). 

Newell Beach, 5 km NE of Mossman, Elix 17430 (CANB). Daintree NP., Mossman Gorge Section, near western 

border of National Park along Mossman creek, Mangold 36 s (F). Mossman Gorge NP., near Mossman, Hale 

831842, 832769 (US). Mt. Lewis Rd. 4 km N from Kennedy Hwy., W of Mossman, Hale 832572, 832590 (US). 

S of Mossman, Weber 227890 (CANB, COLO, US). Kuranda, 1893, Wilson '2' (H-Raes.). End of Clohesy River 

Rd. 16 km SE Kennedy Hwy., W of Cairns, Hale 830610, 830840, 832334 (US). Thompson Rd., Edmonton, 9 

km S of Cairns, Elix 17621 (CANB). Atherton Tablelands: Davies Creek Rd. 17 km S of Kennedy Hwy., S of 

Davies Creek Falls NP., E of Mareeba, Hale 831267, 832604, 832614 (US); Danbulla Forest Drive, 4 km E of 

Tinaroo, Hale 831185 (US); Lake Euramoo, Lumbsch & Mangold 19127 za (F); Lake Barrie NP., Hale 830751 

(US); Lake Eacham NP., Mangold 29 ad (F); Area below crater, Mt. Hypipamee NP., S of Atherton, Hale 

831518, 832524, 832758 (US); 18 km S of Atherton, Mt. Hypipamee, at Crater lake, Tibell 15421, 15432 (UPS); 

Plath Rd. logging head, 9 km W of Plath Rd., off Kennedy Hwy, Herberton Range, S of Atherton, Hale 830981, 

830995, 832574 (US); SW of K-1 tree rd. off Palmerston Hwy., 11 km from main hwy. and 2 km N of 

S.Johnstone Forestry Camp, SE of Millaa Millaa, Hale 831883, 832594, 832680, 832692, 832697 (US); Zillie 

Falls, Lumbsch & Mangold 19141 b (F); Tumoulin Rd., 5 km from turnoff to Ravenshoe, Mangold 30 n, zh (F); 

Tully Falls NP., S of Ravenshoe, Hale 832444 (US). Francis Range, Woopen Creek Rd., 25 km in from Bruce 

Hwy, NW of Innisfail, Hale 830714, 831901, 831951 (US). Dawson logging area, State Forest Reserve 605, 24 

km S of Koombooloomba turnoff, WSW of Tully, Hale 830693, 832196, 832602 (US). State Forest area on 

Tully Rd., 1 km from jct. with S. Mission Beach Rd., S of Mission Beach, Hale 831123, 831198 (US). Cardwell 

Range, Echo & Davidson Creek Drive, 46 km SE of Ravenshoe, Elix 16130 (CANB). Blencoe Creek, Cardwell 

Range, 48 km NW of Cardwell, Elix 20116, 20132 (CANB). Kirrima SF., Cardwell Range, 24 km WNW of 

Cardwell, Elix 15679, 15715 (CANB), Streimann 28535 (B, CANB), 223545 (H, US). Edmund Kennedy NP., 

on Clift Rd., NW of Cardwell, Hale 830832, 832593, 832817 (US). 1 km N of Murray Falls, W of Kennedy, 

Hale 831394 (US). About 7.5 km E of Wallaman Falls, W of Ingham, Hale 831882, 832350, 832591 (US). Little 

Crystal Creek Falls, Mt. Spec NP., the falls on Paluma Rd., Hale 831119 (US). Conway SF., 18 km E of 

Prosperpine, Elix 20778, 20787, 20791 (CANB). Clarke Range, 46 km SSW of Proserpine, Elix 20860 (CANB). 

Eungella NP., Finch Hatton Gorge: Lumbsch & Mangold 19115 h, 19116 a, g, (F); Verdon 5245 (CANB). 

Wooroi Forest, 2 km W of Tewantin: Elix 10423 (CANB); Hale 831877 (US). Noosa Heads, Noosa NP.: A. & 

K. Kalb 25793 (hb. Kalb); Palm Grove Track, Hale 831296, 831760 (US). Mapleton Falls, 13 km W of 

Nambour, Hale 830287 (US). New South Wales: Nightcap NP., Mnt. Nardi/Mnt. Matheson Track, Mangold 22 

p, zi (F). Iluka Nature Reserve, 50 km NE of Grafton: Hale 58728 (US); Mangold 23 k, m, n (F). Dorrigo NP., 

Sassafras Creek Track, Mangold 25 g (F).


Myriotrema subconforme (Nyl.) Hale 

Mycotaxon 11: 135 (1980). Bas.: Thelotrema subconforme Nyl., J. Linn. Soc. London, Bot. 20: 53 (1883). 

Type: Malaya, Malacca, May 1864, Maingay 64 (BM!-lectotype, selected by Hale [1981: 294]; H-Nyl. 22587!-, 

FH-Tuck.!-isolectotypes). 


Fig. 88. Myriotrema subconforme: growth habit (A), ascomata (B), ascospore (C) and ascospore 

showing amyloid reaction (D). A.: BM-lectotype; B.-D.: Mangold 31 l. Bar= A: 1.25 mm; B: 0.5 

mm; C, D: 3 µm. 

Thallus epi- to hypophloedal, moderately thin, up to c. 200 µm high, (pale) olive to (pale) 

grayish-green. Surface shiny, smooth, rugose and/or distinctly verrucose to verruculose, 

unfissured to fissured. True cortex present, continuous, thick to very thick, up to c. 60 µm 

thick, consisting of periclinal hyphae. Algal layer well developed, continuous, becoming 

incontinuous due to crystal inclusions, calcium oxalate crystals abundant, small to large, 

scattered to clustered. Vegetative propagules not seen. Ascomata inconspicuous but often 

very abundant, (very) small, up to c. 200 µm in diam., roundish, apothecioid, solitary to 

slightly fused, immersed. Disc usually not visible from surface, rarely becoming partly 

visible, pale flesh colored, distinctly pruinose. Pores tiny to small, up to c. 50(80) µm in 

diam., predominantly irregular and ±split, proper exciple becoming apically to more rarely 

entirely visible from surface, whitish, shrunken, incurved. Thalline rim margin (moderately) 

small, roundish to somewhat irregular, (moderately) thin, entire to split, whitish or brighter 

than thallus, flush with thallus to somewhat raised, thalline rim incurved. Proper exciple free 

in upper parts, thin, hyaline internally, pale yellowish to yellowish-brown marginally, often 

grayish to more darkened apically, non-amyloid. Hymenium up to c. 80 µm high, non-


inspersed, strongly conglutinated, paraphyses bent, unbranched to slightly branched towards 

the exciple, moderately interwoven, with slightly to distinctly thickened, irregular tips, lateral 

paraphyses and columellar structures absent. Epihymenium predominantly (moderately) 

thick, hyaline, with grayish granules. Asci 8-spored, tholus thick, thin at maturity. Ascospores 

(very) small, submuriform, cell walls thick, endospore (moderately) thick, non-halonate or 

with thin halo in younger stages, hyaline, moderately to strongly amyloid, oblong to ellipsoid 

to more rarely fusi- or claviform, with roundish to subacute ends, loci roundish to angular, 

subglobose to oblong to more often irregular, with hemispherical to conical end cells, 

transverse septae distinct, moderately thin to thick, becoming irregular in older ascospores, 

10-18(20) x 6-9 µm with 4-6 x 1-3(4) loci. Pycnidia not seen. 



subconforme was collected in Australia on tree bark 

in coastal forests, mangroves and monsoon forests 

in low altitudes ranging from sea level to 150 m. It 

is common in the north-western Northern Territory 

and northern Queensland. This is the first report for 

Australia. The paleotropical species was previously 

known from India (Hale, 1981), Sri Lanka (ibid.), 

Malaya, Philippines (ibid.), Java (ibid.), Sarawak 

(ibid.) and Solomon Islands. 

N OTES – Among Myriotrema species with 

±small, immersed ascomata with free proper 

exciple and small, hyaline and amyloid ascospores, 

M. subconforme is distinguished by the thin, 

distinctly corticate thallus with numerous crystal 

inclusions, ascospores with ±irregular, distinctly 

submuriform loci arrangement and the absence of 

secondary compounds. Myriotrema viridialbum and 

M. rugiferum are similar, but can be readily 


M. subconforme. 

distinguished by larger ascomata, distinctly exceeding 200 µm in diam., and their chemistry, 

containing hypoprotocetraric acid (M. viridialbum) and psoromic acid (M. rugiferum), 

respectively. 

SPECIMENS EXAMINED – Australia, Northern Territory: Channell Point, 23 km NNW of Daly River, Elix 

27702 (B, CANB). Queensland: Easternfall of Mc Ilwraith Range, about 15 km upstream from the upper 

crossing at Massy Creek on Silver Plains Stn., Clarkson 2638 (BRI). Cape Flattery sand dunes, McCracken 

694741 (BRI). Hope Vale Cutoff, NE of Cooktown, Hale 830761 (US). Helensvale-Rossville Rd., 29 km S of 

Cooktown, Elix 17386 (B, CANB).Cape Tribulation Area: Track to Cape Tribulation Beach, Mangold 32 n (F); 

Myall Beach, Lumbsch & Mangold 19160 p, 19161 k, t, Mangold 31 l, r (F); 2 km W of main rd. between Oil 

Palms and Coopers Creek, N of Daintree, Hale 831857 (US). Fitzroy Island on Great Barrier Reef, 25 km E of 

Cairns, A. & M. Aptroot 22316 (ABL). Fresh Water Gorge, outside of city of Cairns, Hale 831657 (US). 

Thompson Rd., Edmonton, 9 km S of Cairns, Elix 17607 (B, CANB). Stallion Pocket logging area, 14 km from 

Gillies Hwy. and 1 km E from Mulgrave River Forestry rd., S of Gordonvale, Hale 832398 (US). State Forest 

area on Tully Rd., 1 km from jct. with S. Mission Beach Rd., S of Mission Beach, Hale 831541 (US). Few km N 

of Cardwell, Edmund Kennedy NP., A. & K. Kalb 34234 (hb. Kalb). Edmund Kennedy NP., on Clift Rd., NW of 

Cardwell, Hale 832365, 832393 (US). Hinchinbrook Islands, Rainforest Creek (Bishop Creek), Stevens 689498, 

689499 (BRI). Solomon Isld., Hill 10043 (US).


Myriotrema temperatum Mangold spec. nov. ined. 

Type: Australia, New South Wales, Chaelundi Rd., near Fellabindi Rd., Chaelundi SF., 51 km NW of 

Dorrigo, Streimann 47489 (CANB-holotype). 

ETYMOLOGY – The epithet refers to the temperate distribution of the species. 


Fig. 90. Myriotrema temperatum: growth habit (A), ascomata (B), ascoma and thallus section (C), 

pycnidia (D), condidia (E), ascospores (F) and same ascospores showing amyloid reaction (G). A.: 

Verdon 3882; B.-G.: CANB-holotype. Bar= A: 1.5 mm; B: 1 mm; C: 150 µm; D: 0.1 mm; E: 5 

µm; F, G: 4 µm.


Thallus variable, epi- to hypophloedal, (moderately) thick, up to c. 600 µm high, (pale) 

olive to yellowish-olive, sometimes with bright speckles. Surface dull to slightly shiny, 

smooth, ±rugose, usually distinctly to strongly verrucose to verruculose, unfissured to more 

often distinctly fissured. Cortex structures variable, often true cortex present, ±continuous, 

thin, hyaline to slightly yellowish, up to c. 25 µm thick, consisting of periclinal to irregular 

hyphae, sometimes lacking a true cortex, then covered by a thin ±incontinuous protocortex. 

Algal layer well developed, continuous, calcium oxalate crystals abundant, often large, 

clustered, sometimes forming column-like structures. Vegetative propagules not seen. 

Ascomata variable, sparse to abundant, becoming moderately large, up to c. 600 µm in diam., 

roundish, apothecioid, solitary to marginally or more rarely entirely fused, immersed to 

emergent, then usually ±conical. Disc often becoming partly visible, pale flesh colored, 

epruinose. Pores (moderately) small, sometimes becoming wide, up to c. 400 µm in diam., 

predominantly roundish, entire, proper exciple not visible from surface. Thalline rim margin 

moderately thick, distinctly bright, off-white or brighter than thallus, in specimen with 

predominantly immersed ascomata sometimes sunken, thalline rim incurved. Proper exciple 

fused to apically slightly detached, moderately thin to moderately thick, hyaline internally, 

grayish to pale grayish-brown marginally, non-amyloid. Hymenium up to c. 100 µm high, 

non-inspersed, moderately to weakly conglutinated, paraphyses straight, slightly interwoven, 

with slightly to distinctly thickened tips, lateral paraphyses lacking and columellar structures 

absent. Epihymenium indistinct to thin, hyaline, rarely with fine grayish granules. Asci 8spored, 

tholus moderately thick, thin when mature. Ascospores (very) small, transversely 

septate, cell walls and endospore (moderately) thick, usually with a thick halo, hyaline, 

distinctly to strongly amyloid, oblong to ellipsoid or somewhat fusi- to rarely claviform, with 

roundish to subacute ends, loci roundish, oblong to lentiform, with hemispherical to conical 

end cells, septae (moderately) thin, regular, 10-17 x 5-8 µm with 3-4(5) loci. Pycnidia present, 

immersed or in thallus warts, with bright rimmed, dark pore, conidia fusiform, up to c. 5 x 2 

µm. 


and subpsoromic (traces) acids. 


temperatum grows on tree bark in subtropical 


It is moderately common in the Queensland/New 

South Wales border region and northern New South 

Wales and currently only known from there. 


predominantly corticate thallus with a rugose, 

verrucose to verruculose surface, abundant calcium 

oxalate crystals, ascomata with a fused proper 

exciple and a distinct, entire bright margin, small, 

transversely septate, hyaline, amyloid ascospores 

with a distinct halo, and the presence of the 

psoromic acid chemosydrome. It may be difficult to 

distinguish from M. microporum and M. clandestinum, 

but it differs from both in the thallus 

surface structure and the thicker thalline rim 

margin. Further differences to M. microporum are 

listed under this species. Myriotrema clandestinum 


M. temperatum.


differs further by having longer ascospores (up to 25 µm) that either lack a halo or have only 

a thin halo. Another similar species is M. glaucophaenum, which can be readily distinguished 

by larger ascomata with a free proper exciple and an often split to lacerate thalline rim 

margin. 

SPECIMENS EXAMINED – Australia, Queensland: Cunninghams Gap NP., 50 km NE of Warwick, Hale 

831346 (US). Lamington NP.: Python Rock Track, Hale 831011 (US); Beechmont Range, Binna Burra, A. & K. 

Kalb 19864, 21581 (hb. Kalb). New South Wales: Mt. Warning NP., W of Murwillumbah, Hale 832565 (US). 

Trail from Black Butt Picnic Area, Wiangaree Forest Drive, NE of Kyogle, Hale 831732 (US). Gibbergunyah 

Roadside Reserve, Nightcap Forest Drive, Whian Whian SF., W of Mullumbimby, Hale 831476 (US). 37 km N 

of Ebor, Chaelundi Mt., Verdon 3882 (CANB, H, TNS). Dorrigo NP., summit of Dorrigo Mt., Elix 3480 

(CANB). Beech Plateau, Werrikimbe NP., 80 km NW of Port Macquarie, Streimann 63985 (B). 

Myriotrema trypaneoides (Nyl.) Hale 

Mycotaxon 11: 135 (1980). Bas.: Thelotrema trypaneoides Nyl., Ann. Sci. Nat. Bot. ser. 4, 19: 335 (1863). 

Leptotrema trypaneoides (Nyl.) Riddle, Bull. Torrey Bot. Cl. 43: 151 (1916). Type: Cuba, Wright 156 (FH- 

Tuck.–lectotype selected by Hale [1978: 54]; US!–isolectotype). 

Thelotrema subterebrans Nyl., Flora 59: 561 (1876). Leptotrema subterebrans (Nyl.) Zahlbr., Cat. Lich. 

Univ. II: 640 (1923). [fide Hale 1978]. 

Thelotrema laevius Vain., Journ. of Botan. 34: 207 (1896). Leptotrema laevius (Vain.) Zahlbr., Cat. Lich. 

Univ. II: 635 (1923). Type: St. Vincent, Elliott s.n. (TUR-Vain. 26774!-lectotype, here selected). 


Thallus epi- to hypophloedal, (moderately) thick, up to c. 1 mm high, sometimes bulging 

and splitting away from the substrate, (pale) greenish gray, often with a grainy-speckled 

pattern. Surface ±shiny, smooth, continuous to verruculose, unfissured to rarely somewhat 

rimose. True cortex present, continuous, up to c. 40µm thick, hyaline, predominantly 

consisting of periclinal hyphae. Algal layer continuous and well developed, calcium oxalate 

crystals usually abundant, small to large, predominantly clustered, often forming columnar 

structures. Vegetative propagules not seen. Ascomata inconspicuous, (moderately) small, up 

to c. 500 µm in diam., roundish, predominantly perithecioid, solitary, immersed to slightly 

emergent, then hemispherical. Disc not visible from surface. Pores tiny, up to c. 50 µm in 

diam., roundish, entire, apical proper exciple becoming visible from surface, forming a fused 

inner pore margin, pore margin level thallus to often sunken, incurved, bright-translucent to 

(dark-)gray. Thalline rim margin moderately thick, roundish, entire, concolorous with thallus, 

thalline rim not developed or not distinguishable from surface, in emergent ascomata 

incurved. Proper exciple fused, (moderately) thin, pale yellowish internally, yellowish- to 

orange-brown marginally, sometimes dark-brown apically, non-amyloid. Hymenium up to c. 

250 µm high, densely inspersed, moderately conglutinated, paraphyses thin, straight to 

slightly bent, parallel, unbranched, tips unthickened to slightly thickened, lateral paraphyses 

and columellar structures absent. Epihymenium indistinct to moderately thin in older stages, 

hyaline to yellowish or orange towards proper exciple, without granules or crystals. Asci 8spored, 

tholus moderately thick, thin when mature. Ascospores (moderately) small, 

eumuriform, cell walls and endospore (moderately) thick in younger stages, moderately thick 

to moderately thin at maturity, becoming brown in early stages, non-amyloid, oblong to 

ellipsoid with roundish to narrowed-roundish ends, loci large in early stages, becoming small 

with late maturity, roundish to slightly angular, subglobose to oblong or irregular with same 

shaped end cells, transverse septae (moderately) thin, ±regular, 25-40 x 8-15 µm with 6-12 x 

1-6 loci. Pycnidia not seen.


Fig. 92. Myriotrema trypaneoides: growth habit (A), ascomata (B), ascoma and thallus section (C), 

ascospore (D), young ascospore (E) and old ascospore (F). A.: Hale 831360; B., C.: TURholotype 

of T. laevius; D.-F.: Lumbsch & Mangold 19167 v. Bar= A: 1 mm; B: 0.15 mm; C: 225 

µm; D: 7 µm; E: 4 µm; F: 6 µm. 

CHEMISTRY – Thallus K+yellow, C-, PD+orange; containing constictic and stictic acids 

(majors), cryptostictic, hypoconstictic and hypostictic (minors) acids. 

ECOLOGY AND DISTRIBUTION – Myriotrema trypaneoides was collected in Australia on tree 

bark in tropical rainforests in altitudes ranging from sea level to 700 m. It is moderately 

common in northern and central Queensland. This is the first report for Australia. This 

pantropical species has been recorded from Central America, India (Patwardhan & Kulkarni, 

1977) and the Philippines (ibid.). 

NOTES – This taxon is characterized by the shiny, corticate, thick thallus, the immersed to 

slightly emergent perithecia with tiny pores, the dark exciple, an inspersed hymenium, the 

moderately small, brown, muriform ascospores with a thick-walled young stage and the 

presence of the stictic acid chemosydrome. It is similar to M. desquamans, but can be readily 

distinguished by the darker proper exciple and the inspersed hymenium. The only similar


species with inspersed hymenium is L. bisporum 

from Guadeloupe, which differs in having larger 

ascospores up to 180 µm long. 

SPECIMENS EXAMINED – Australia, Queensland: Cape 

York Peninsula, Iron Range NP., Hale 830051, 830061 (US). 

Near Cedar Bay NP., Rd. to Cooktown, Mangold 34 x (F). 

Cape Tribulation Area: Cape Kimberley, Lumbsch & 

Mangold 19167 v (F); Cape Tribulation NP., Hale 69193, 

831360 (US). Fresh Water Gorge, outside of city of Cairns, 

Hale 831731 (US). Fitzroy Island on Great Barrier Reef, A. 

& M. Aptroot 22315 (ABL). Crystal Cascades, Lumbsch & 

Mangold 19129 k (F). Atherton Tablelands: Lake Tinaroo, 

Lumbsch & Mangold 19127 b, zb (F); Curtain Fig Tree, 

Lumbsch & Mangold 19128 o (F); Souita Falls, Lumbsch & 

Mangold 19155 j, 19156 e (F); S of Tolga on Kennedy Hwy, 

Hale 831617 (US); Wongabel SF., Hale 830604 (US); 27 km 

on Mulgrave River Rd., SW of Gordonvale, Hale 830313, 

830681 (US). Francis Range, Woopen Creek Rd., Hale 

832067 (US). Murray Falls, W of Kennedy, Hale 831399 Fig. 93. Australian distribution of 

(US); 2 km N of Murray Falls, W of Kennedy, Hale 831244, M. trypaneoides. 

832323 (US). Conway Peninsula, E of Proserpine, Scott 620 

(BRI). Conway SF., 18 km E of Proserpine, Elix 20185, 

20202 (CANB). N of Emu Park, near Rockhampton, Selling 16442 (S). 

Myriotrema viridialbum (Kremp.) Hale 

Mycotaxon 11: 135 (1980). Bas.: Thelotrema viridialbum Kremp., Flora 49: 221 (1876). Ocellularia 

viridialba (Kremp.) Müll. Arg., Flora 70: 398 (1887). Rhabdodiscus viridialbus (Kremp.) Vain., Ann. Acad. 

Scient. Fenn. A15(6): 184 (1921). Type: Brazil, Rio de Janeiro, Glaziou 3193 (M-lectotype, selected by Hale 

[1972 in herb.]; C!-isolectotype). 

Thelotrema myrioporoides Müll. Arg., Bull. Soc. Bot. Belgique 32: 147 (1893). Myriotrema myrioporoides 

(Müll. Arg.) Hale, Mycotaxon 11: 134 (1980). Type: Costa Rica, Boruca, 1893, Tonduz s.n. (com. Pittier 6104) 

(G!-lectotype, selected by Hale [1978: 45]). 

Thelotrema leucohymenium Zahlbr., Denkschr. math.-naturw. Classe K. Akad. Wiss. Wien 83: 120 (1909). 

Type: Brazil, Sao Paulo, 16.07.1901, Schiffner s.n. (W-holotype, BM!-isotype). 

Thelotrema steyermarkii Hale, Phytologia 27: 496 (1974). Myriotrema steyermarkii (Hale) Hale, Mycotaxon 

11: 135 (1980). Type: Venezuela, Bolivar, Steyermark 98008 (US!-holotype). 


Thallus corticolous, sometimes overgrowing adjacent bryophytes, predominantly epi- to 

sometimes slightly hypophloedal, thick, up to c. 700 µm high, pale yellowish- to greenish 

gray or (pale) olive. Surface dull to shiny, smooth, continuous to rugose to more rarely 

verrucose, unfissured to more often distinctly fissured. True cortex present, ±continuous, 

sometimes yellowish, up to c. 50 µm thick, predominantly consisting of periclinal hyphae. 

Algal layer well developed, continuous, calcium oxalate crystals lacking or sparse, then large 

and clustered, distinct medullar layer present. Vegetative propagules not seen. Ascomata 

variable, small to large, up to c. (500)800 µm in diam., predominantly roundish, apothecioid, 

solitary to often fused, immersed to ±distinctly emergent in older ascomata, then 

hemispherical to urceolate. Disc sometimes becoming partly visible from surface, pale flesh 

colored, pruinose. Pores small to gaping, up to c. (300)600 µm in diam., roundish, entire to 

slightly split, proper exciple often becoming apically or entirely visible from surface, offwhite, 

sometimes slightly shrunken, incurved. Thalline rim margin becoming moderately 

wide to gaping, roundish, moderately thick, predominantly ±entire, whitish or brighter than 

thallus, thalline rim incurved. Proper exciple fused to partly or entirely free, moderately thick


Fig. 94. Myriotrema viridialbum: growth habit (A, B), ascomata (C), pycnidia (D), conidium (E), 

ascospores (F) and ascospores showing amyloid reaction (G). A.: US-holotype of T. steyermarkii; 

B.: Hale 830899; C.-G.: C-isolectotype. Bar= A: 1 mm; B: 1.5 mm; C: 0.5 mm; D: 0.2 mm; E: 2 

µm; F, G: 7.5 µm. 

to moderately thin, with thin or absent hyaline area internally, entirely or marginally grayish, 

sometimes faintly amyloid (reddish). Hymenium up to c. 160 µm high, non-inspersed, 

sometimes interspersed with large, ±columnar aggregates of bacilliform crystals (dissolving 

in KOH) strongly conglutinated, paraphyses thick, irregular and often distinctly septated, 

±interwoven, with thickened, irregular tips, lateral paraphyses and true columella absent, 

columella-like structures sometimes present in fused ascomata. Epihymenium moderately 

thick, hyaline, with grayish granules. Asci 8-spored, tholus moderately thick to moderately 

thin. Ascospores (very) small, predominantly submuriform, cell walls and endospore 

(moderately) thick, often covered by a thin halo, hyaline, distinctly to strongly amyloid, 

subglobose to ellipsoid or somewhat claviform, with roundish to subacute, rarely acute ends, 

loci roundish, subglobose to lentiform, with same shaped to more hemispherical or rarely 

conical end cells, septae (moderately) thick, often irregular, 10-20 x 5-12 µm with 3-6 x 1- 

2(3) loci. Pycnidia often present, variable, immersed or in distinct thallus warts, with


brownish pore area surrounded by a ±eroded, bright region, conidia fusiform, up to c. 6 x 1 

µm. 

CHEMISTRY – Thallus K-, C-, PD-, UV±; containing hypoprotocetraric (major), 4-0demethylnotatic, 

convirensic, conhypoprotocetraric (minors to traces) and conprotocetraric 

(trace) acids, and sometimes lichexanthone. 


viridalbum occurs on tree bark in tropical 

rainforests at elevations ranging from 600 to 

1100 m altitude. It is rare in Australia, occurring 

only in northern Queensland. This is the first report 

for Australia. Previously only known from the 

Neotropics. 

NOTES – The species is characterized by a thick, 

epiphloedal, corticate thallus, small, submuriform, 

amyloid ascospores with distinctly thickened parts 

and the presence of the hypoprotocetraric acid 

chemosydrome. Myriotrema glaucophaenum is 

similar, but can be readily distinguished by 

transversely septate ascospores and psoromic acid 

(for a discussion of the columella-like structures in 

the 'M. viridialbum-group' see under M. glaucophaenum). 

Also similar is M. microporum, which 

differs in having transversely septate ascospores 

and psoromic acid. Myriotrema rugiferum is 


M. viridialbum. 

another taxon that can be confused with M. viridialbum, but differs by smaller ascomata (not 

exceeding 400 µm in diameter) with less widened pores. The Australian collections agree 

well with the Neotropical specimen, except for the absence of lichexanthone and smaller 

ascomata. An unusual feature in M. viridialbum is the presence of large crystal aggregates in 

the hymenium, found in the type and in the type collection of T. leucohymenium (the epithet 

even refers to the phenomenon) and T. myrioporoides. It only occurs in older samples. 

SPECIMENS EXAMINED – Australia, Queensland: Mt.Windsor, 5 km W of new Forestry Camp, NW of 

Mossman, Hale 830663, 830899 (US). Cardwell Range: Blencoe Creek, 48 km NW of Cardwell, Elix 20134, 

20138 (CANB); 24km WNW of Cardwell, Streimann 28545 (B, CANB). Lannercost SF., Blue Water Creek, 

Old Mill Rd., 39 km WSW of Ingham, Elix & Streimann 15572 (CANB). Venezuela, Paramo La Negra, Estado 

Merida, (as T. steyermarkii, Lichenes Americani Exsiccati no. 199), Hale 42871 (MEL). Costa Rica, Cartago, 

Cordillera de Talmanaca, Filson 16272 (MEL). 

2. 9. 8. Nadvornikia Tibell, Beih. Nova Hedw. 79: 672 (1984). Type species: Acolium 

hawaiense Tuck. [=N. hawaiensis (Tuck.) Tibell]. 

Stephanophoron Nádv., Ann. Mycol. 40: 136 (1942), nom. illeg. [non Stepahnophoron Flotow]. 

Nadvornikia hawaiensis is the only known species in Australia, for a description see there. 

NOTES – This genus was introduced by Tibell (1984) in the family Caliciaceae with N. 

hawaiensis as the only member. Tuckerman (1867) already pointed out the similarities to 

Thelotremataceae in his original description of the species (as Acolium hawaiense). Harris


(1990) proposed to classify the genus in this family, which was later accepted by Tibell 

(1996) and has been recently confirmed using molecular analysis (Lumbsch & al., 2005). 

Two additional species were since described for the genus, N. diplotylia (Pant & Awasthi, 

1989), which is included as a synonym in N. hawaiensis [following Tibell (1996)], and N. 

sorediata (Harris, 1990). The latter species (only known from the Neotropics) differs from N. 

hawaiensis by the presence of soralia. 

The genus is readily distinguished by the mazaedious ascomata with small, brown 

bilocular ascospores and no other similar genus is known. 


Nadvornikia hawaiensis (Tuck.) Tibell 

Beih. Nova Hedwigia 79: 672 (1984). Bas.: Acolium hawaiense Tuck., Proc. Amer. Acad. Arts Sci. 7: 232 

(1866). Type: Hawaii, Oahu, Wailaua Mts., Mann s.n. (UPS!-lectotype, here selected [annotated as 'syntype', 

Tibell 1987 - see also notes]). 

Tylophoron diplotylium Nyl., Bull. Soc. Linn. Normandie 2: 46 (1868). Nadvornikia diplotylia (Nyl.) Pant & 

Awasthi, Biovigyanam 15(1): 12 (1989). Type: New Caledonia, Wagap, 1863, ("D23"), s.c. (H-Nyl. 40422!holotype). 


Fig. 96. Nadvornikia hawaiensis: growth habit (A), ascoma (B), ascoma section (C), ascospores (D, 

E). A.: H-lectotype of T. diplotylium; B., D., E.: Tibell 12673; C.: Mangold 36 x. Bar= A: 3 mm; 

B: 0.5 mm; C: 300 µm; D: 10 µm; E: 3 µm.


Thallus epi- to hypophloedal, moderately thick, up to c. 300 µm high, often bulging and 

flaking away from the substrate, pale greenish- to pale yellowish-gray. Surface dull to 

glittering or slightly shiny, smooth to somewhat roughened, ±rugose, continuous to usually 

distinctly to strongly verruculose, unfissured. Cortex structures absent or with a thin, 

incontinuous protocortex up to c. 20 µm thick. Algal layer moderately well developed, 

continuous to incontinuous, calcium oxalate moderately abundant, predominantly small, 

scattered to clustered. Vegetative propagules not seen. Ascomata conspicuous, large, up to c. 

1.5 mm in diam., roundish to slightly irregular, mazaedious, solitary, strongly emergent, 

subglobose to urceolate in older ascomata. Pores becoming wide to gaping, up to c. 800 µm in 

diam., roundish to somewhat elongated, entire to slightly split, apical proper exciple often 

becoming visible from surface due to eroded thalline rim, forming a brownish to reddishbrown 

ring, and/or the proper exciple becomes visible as a distinctly raised, slightly incurved 

to erect, corona-like velum, slightly pruinose. Thalline rim margin thick, roundish to slightly 

elongate, becoming distinctly eroded with age, brighter then thallus, thalline rim becoming 

erect, with same surface as thallus. Proper exciple fused, in older ascomata becoming apically 

free, thick, hyaline basally, pale orange to orange-brown apically, in upper, internal parts 

often covered by brownish-gray granules, non-amyloid. Hymenium only visible in younger 

ascomata, clear, distinctly conglutinated, paraphyses thin, slightly interwoven, unbranched, 

straight to somewhat bent, in mature stages distinctly mazaedious, lateral paraphyses and 

columellar structures absent. Asci 8-spored, tholus not visible. Ascospores very small, bilocular, 

with thickened cell walls and septae, non-halonate, surface becoming distinctly, 

±irregularly ornamented with age, brown, non-amyloid, predominantly oblong to fusiform or 

more rarely subglobose, with roundish to somewhat subacute ends, loci predominantly 

hemispherical, 6-10 x 4-6 µm. Pycnidia not seen. 


constictic (majors), hypostictic (minor to trace), α-acetylconstictic and hypoconstictic (traces) 

acids. (See notes regarding the presence of lichexanthone.) 

ECOLOGY AND DISTRIBUTION – Nadvornikia 

hawaiensis was collected in Australia on tree bark 

in (sub)tropical rainforests, rarely in wet 

sclerophyll forests, in altitudes ranging from 50 to 

900 m. It is moderately common and widespread in 

Queensland and in the Queensland/New South 

Wales border region. It is pantropical, being 

recorded from Hawaii, Brazil (Tibell, 1984), the 

Andman Islands (Pant & Awasthi, 1989) and New 

Caledonia. 

N OTES – This taxon is unusual in 

thelotremataceaen Graphidaceae for having 

mazaedious ascomata. Nadvornikia includes two 

species, the second being N. sorediata from the 

Neotropics. The phylogenetic placement of the 

genus in Thelotremataceae was confirmed using 

molecular analysis (Lumbsch & al., 2004), see also 

Tibell (1984) and Harris (1990) for a more detailed 

discussion on the taxonomy of the genus. The 


N. hawaiensis. 

syntypes in FH were not available for study, Tibell (1989) neither saw this material but a 

specimen in UPS. Harris (1990) states that he saw two isotypes but does not give any further


information. Since the UPS collection is the only widely available type material it is here 

selected as lectotype. The chemistry of this species caused some confusion. Tibell (1984, 

1989) stated the UV-reaction ('brilliant UV+ yellow' and 'faintly greenish green' in the 

Australian material; 'intense UV+ fluorescent') and concluded the presence of lichexanthone, 

additionally, he notes about the Australian material that "no secondary products have been 

identified". Consequently Pant & Awasthi (1989) classified Nadvornikia diplotylia (ex 

Tylophoron diplotylium) as a distinct species (formerly treated as a synonym by Tibell), based 

on the lack of lichexanthone and the presence of the stictic acid chemosydrome. The chemical 

analysis of the examined specimen conducted here confirmed Harris' (1990) statement (type 

collection and several other specimen tested) that they contain the stictic acid chemosydrome 

but lack lichexanthone. 

SPECIMENS EXAMINED – Australia, Queensland: Big Tableland, 26 km S of Cooktown, Elix 17265 (CANB). 

Daintree NP., Mossman Gorge Section, near western border of National Park along Mossman creek, Mangold 36 

w, x (F). Babinda Boulders, A. & M. Aptroot 22391 (ABL), Lumbsch & Guderley 11151 e (F). Near Yabba Rd., 

6 km N of Jimna, Rogers 2247 (BRI). Mt. Glorious, 2 km W of Maiala picnic area, Rogers 2577 (BRI). B.Rose 

Park, Samford-Mt.Glorious Rd., Rogers 2702, 2703 (BRI). Duck Creek rd. near Lamington NP., close to 

O'Reilly's, (Lichenoth. Graecensis 12), Mayrhofer, Hierzer & Rogers s.n. (M). Lamington NP.: 13 km SW of 

Beechmont, along Blue Pool Track, (Calic. exs. 121), Tibell 12673 (F, COLO); Binna Burra, Aptroot 46103 

(ABL). Springbrook-Numinbah-Nerang intersection, Rogers 2975 (BRI). New South Wales: Unumgar SF., 4 km 

S from Qld. border, Lumbsch & Mangold 19176 q (F). 

2. 9. 9. Pseudoramonia Kantvilas & Vezda, Lichenologist 32: 344 (2000). Type species: 

Pseudoramonia. stipitata (Vezda & Hertel) Kantvilas & Vezda. Type: Venezuela, Estado 

Merida, Sierra de Santo Domingo, Hertel & Oberwinkler 10412 (hb. Hertel-holotype, hb. 

Vezda-isotype). 

Pseudoramonia richeae is the only known species in Australia, for a description see there. 

NOTES – This recently introduced genus (Kantvilas & Vezda, 2000) consists of two 

species, P. richeae which is so far only known from Tasmania and the neotropical P. stipitata 

(for distinguishing characters of the two taxa see under P. richeae), that differ from other 

thelotrematoid taxa by distinctly stipitate ascomata. Otherwise Pseudoramonia is similar to 

Topeliopsis, Melanotopelia and the ‘Leptotrema schizoloma-group’. 


Pseudoramonia richeae Kantvilas & Vezda 

Lichenologist 32: 344 (2000). Type: Tasmania, Mt.King William, Kantvilas 105/84 (HO!-holotype, hb. 

Vezda-isotype). 


Thallus corticolous to foliicolous epi- to endosubstratic, moderately thin, up to c. 200 µm 

high, pale gray to yellowish-gray or rarely greenish-gray. Surface dull to slightly shiny, 

smooth, continuous, unfissured to slightly fissured. Cortex structures variable, predominantly 

covered by an incontinuous protocortex up to c. 25 µm thick, in ascoma region often 

becoming distinctly conglutinated, forming a true cortex of irregular to periclinal hyphae.


Fig. 98. Pseudoramonia richeae: growth habit (A, B), ascomata (C, D), ascomata sections (E. F), 

hymenium and asci (G) and ascospores (H). A.-H.: HO-holotype. Bar= A: 4 mm; B: 1 mm; C: 0.7 

mm; D: 0.5 mm; E: 150 µm; F: 120 µm; G: 10 µm; H: 12 µm. 

Algal layer poorly developed, incontinuous, calcium oxalate crystals absent. Isidia-like 

structures often present, probably representing immature ascomata (see notes). Ascomata very 

characteristic, conspicuous, moderately small to moderately large, up to c. 600 µm in diam., 

roundish to irregular in fused ascomata, peri- to apothecioid, solitary to fused, stipitate, 

growing successively, either depressed, forming a broad, verrucose base or emerging, forming 

somewhat coralloid structures. Disc usually not visible from surface, rarely becoming 

somewhat visible, grayish, epruinose. Pores small to moderately wide, up to c. 400 µm in 

diam., predominantly irregular or slightly elongate, margin ±split, incurved and often 

distinctly sunken, concolorous with thallus or darkened due to protuberant proper exciple, 

otherwise proper exciple not visible from surface. Thalline rim moderately thick, concolorous 

to somewhat darkened, predominantly depressed-subglobose to irregular with ±strongly 

restricted base, base stiped, stipes up to c. 2 mm long, concolorous with thallus to reddishbrown. 

Proper exciple fused, moderately thick, pale yellowish to pale yellowish-brown 

internally, brown to carbonized marginally, not separated from subhymenium, non-amyloid. 

Hymenium up to c. 90 µm high, non-inspersed, strongly conglutinated, paraphyses straight,


parallel, with unthickened tips, lateral paraphyses indistinct, up to c. 15 µm long, not clearly 

separated from the exciple, true columella absent, columella-like structures sometimes present 

in fused ascomata. Subhymenium conspicuous, usually dark-brown to carbonized in upper 

part, sometimes followed by a lower layer of ±hyaline, loosely organized to 

paraplectenchymatous hyphae, resembling a newly developing hymenial layer. Epihymenium 

indistinct, hyaline and without granules. Asci 8-spored, tholus moderately thin, not visible at 

maturity. Ascospores small, transversely septate, cell walls and endospore thin, distinctly 

halonate, hyaline, non-amyloid, oblong to somewhat fusi- to claviform, with roundish to 

narrowed-roundish ends, loci roundish to slightly angular, subglobose to oblong to slightly 

irregular, with same shaped or hemispherical to conical end cells, septae moderately thin to 

moderately thick, slightly irregular, 12-26 x 3-6 µm with 8-12 loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellowish-brown, C-, PD+ orange-red; containing 

succinprotocetraric (major), protocetraric and fumarprotocetraric (major to minor) acids. 

ECOLOGY AND DISTRIBUTION – Pseudoramonia 

richeae occurs on bark and dead leafs of Richea 

scoparia, in (sub-)alpine heathland shrubs in 

altitudes ranging from 700 to 1300 m. It is a 

moderately common species occurring in several 

regions of Tasmania. 

NOTES – This taxon is characteristic by having 

stipitate ascomata. One other species is known in 

the genus, P. stipitata from Venezuela, which 

differs in having the stictic acid chemosyndrome 

and smaller ascospores. Regarding the presence of 

isidia, the original description (Kantvilas & Vezda, 

2000) is followed here, in which the isidia-like 

structures are defined as immature ascomata. In 

contrast to Kantvilas and Vezda (2000) I found the 

ascomata of P. richeae being carbonized. 

SPECIMENS EXAMINED – Australia, Tasmania: Mount Field 

NP., near Lake Dobson, 65 km WNW of Hobart, Aptroot 

23397 (ABL). 


P. richeae. 

2. 9. 10. Reimnitzia Kalb, Mycotaxon 79: 325 (2001). Type species: Reimnitzia santensis 

(Tuck.) Kalb. 

Reimnitzia santensis is the only known species in the genus, for a description see there. 

NOTES – This monotypic genus was erected (Kalb, 2001) based mainly on the unusual 

structure of the apical hymenium with paraphyses having branched and interwoven tips. Since 

similar structures could be also found in Chapsa, Frisch (2006) points out several 

distinguishing characters to justify its status as a separate genus. Hence Reimnitzia is 

characterized by chroodiscoid ascomata and a ‘Thelotrema-type’ exciple and distinguished, 

particularly from Chapsa, by thick-walled ascospores at early stages, ascus tips with a 

“distinct key-hole appearance” (ibid.: 271) (not found in all specimen), “rather lax and less 

distinct” (ibid.: 272) paraphyses, a thallus with large, columnar calcium oxalate crystals and


typical pycnidia that occur in ±strongly emergent thallus warts and on the tips of isidia-like 

structures, and bacilliform conidia. In my studies, I could not confirm the presence of a 

‘Thelotrema-type’ exciple in Reimnitzia. The examined material, which did not include the 

type, however, lacked lateral paraphyses. This agrees well with the protologue of Reimnitzia 

in which the genus is separated from Thelotrema “by the lack of distinct periphyses” (Kalb, 

2001: 326). The genus is accepted here tentatively and needs evaluation by molecular data. 


Reimnitzia santensis (Tuck.) Kalb 

Mycotaxon 79: 325 (2001). Bas.: Thelotrema santense Tuck., Proc. Amer. Acad. Arts Sci. 5: 406 (1862). 

Leptotrema santense (Tuck.) Zahlbr., Cat. Lich. Univ. 2(4): 635 (1923). Type: U.S.A., South Carolina, Santee, 

1860, Ravenel s.n. (BM-lectotype, selected by Salisbury [1972b: 288]). 

(?) Thelotrema heterosporum C. Knight in Bailey, Synops. Queensland Flora, Suppl. 1: 72 (1886) - fide 

Müller 1887 [as synonym of L. mastoideum]. Leptotrema heterosporum (C.Knight) Zahlbr., Cat. Lich. Univ. 

2(4): 635 (1923). Type: Australia, S. Queensland, on moss, C. Knight s.n. (?WELT, not seen - see also notes). 

Leptotrema mastoideum Müll.Arg., Flora 70: 400 (1887) - ?nom. superfl. pro T. heterosporum (if 

conspecific). Type: Paraguay, Balansa 38 (G-holotype). 


Thallus corticolous to rarely muscicolous, epi- to hyposubstratic, moderately thick, up to c. 

300 µm high, rarely bulging, grayish to pale grayish-green or grayish-olive, often with 

distinctly speckled pattern. Surface dull, smooth to mealy to slightly pruinose or rough, 

continuous to verruculose, unfissured to fissured or coarsely cracked. Cortex structures 

absent. Algal layer generally continuous and well developed, appearing incontinuous due to 

crystal inclusions, calcium oxalate crystals abundant, small to large, scattered or clustered, 

forming column-like structures, distinct medulla layer mostly absent, basal thallus often 

formed of conglutinated ±periclinal hyphae, forming a lower cortex-like structure. Isidia often 

present, abundant, concolorous with thallus, unbranched, first globular, becoming vermiform 

with a ±constricted base, up to c. 1.5 mm long. Ascomata conspicuous, up to c. 2 mm in 

diam., roundish to irregular, in particular in fused ascomata, chroodiscoid, solitary to fused, 

erumpent, immersed to slightly raised. Disc usually becoming entirely visible from surface, 

(dark) grayish, pruinose. Proper exciple not visible from surface, thalline rim margin 

becoming wide to gaping, up to c. 1.8 mm in diam., entire to more often ±coarsely split and 

lobed, ±eroded, whitish or brighter than thallus, thalline rim first incurved and moderately 

thin, becoming thick, predominantly erect to slightly recurved in older stages. Exciple fused, 

hyaline to pale yellowish-brown internally, brownish marginally, apically dark-brown to 

rarely slightly carbonized and with grayish granules, non-amyloid. Hymenium up to c. 120 

µm high, non-inspersed, distinctly conglutinated, paraphyses straight, ±interwoven, 

sometimes slightly branched, particularly towards the margins and the apical hymenium, tips 

slightly thickened and irregular, lateral paraphyses not seen, [according to Frisch (2006) 

present, scattered and free, up to 15 µm long], true columella absent, in fused ascomata 

sometimes columella-like structures present. Epihymenium moderately thick, hyaline to 

sometimes brownish marginally, with pale grayish-brown granules. Asci 8-spored, tholus 

moderately thin, thin at maturity. Ascospores small, submuriform, cell walls and endospore 

moderately thick, non-halonate, brown, non-amyloid to rarely very faintly amyloid, 

subglobose to ellipsoid to fusiform, with roundish to subacute ends, loci roundish to angular, 

predominantly irregular, septae (moderately) thin, predominantly irregular, often with a


Fig. 100. Reimnitzia santensis: growth habit (A, B), ascus (C), ascospores (D-F), conidia (G). A., D.- 

F.: Tucker 7614; B.: A. & K. Kalb 30585; C.: Calkin 146; G.: Tucker 32240. Bar= A, B: 2 mm; C: 

10 µm; D-G: 5 µm. 

single, more distinct central septum, 12-25 x 8-12 µm with 4-6 x 1-3 loci. Pycnidia present, 

immersed or in ±globular warts, with brownish to blackish pore area, conidia bacilliform to 

oblong-fusiform, up to c. 6(10) x 1 µm. 

CHEMISTRY – Thallus K-, C-, PD-; no secondary substances detectable by TLC. 

ECOLOGY AND DISTRIBUTION – Reimnitzia santensis occurs on tree bark and in Australia is 

restricted to monsoon forests at low altitudes. It is rare in the area, only known from northwestern 

Northern Territory, and probably southern Queensland (19th century collections). 

(The location in southern Queensland is not marked in the distribution map Fig. 101 due the


uncertain locality). This is a pantropical species that 

has been recorded from the Neotropics and Africa 

(Frisch, 2006). 


isidiate, dull, ecorticate thallus, large, chroodiscoid 

ascomata with pruinose, grayish discs, a fused 

proper exciple, small, brown, submuriform 

ascospores with irregular loci arrangement and the 

absence of secondary compounds. The only similar 

species in Australia is Leucodecton glaucescens, 

for differences see under this species. Thelotrema 

leiospodium Nyl. from Portugal is similar but 

distinguished by a free proper exciple and 

apothecioid rather than chroodiscoid ascomata with 

distinct lateral paraphyses. In contrast to the 

descriptions given by Kalb (2001) and Frisch 

(2006), lateral paraphyses could not be found in the 

examined material. Although the hyaline, inner 

parts of the proper exciple sometimes are formed by 


R. santensis. 

a short-celled plectenchyma that remotely appears to be laterally organized, however, 

distinctly filamentous, paraphyses-like structures were absent. Several characters indicate a 

close relationship to Leptotrema wightii, in particular the habitus, and certain features of 

ascomata, asci and ascospore morphology . The Knight collection from Southern Queensland 

was not available from WELT. Hence the conspecificity to R. santensis is uncertain and it is 

suggested to treat the name as a tentative synonym based on Müller's observations. 

SPECIMENS EXAMINED – Australia, Northern Territory: Kakadu NP., "Gungarre Monsoon Forest", near South 

Alligator, K. & A. Kalb 30585 (CANB, hb. Kalb). U.S.A.: Louisiana: East Baton Rouge Parish, S. Tucker 32240 

(WIS); Livingston Parish, S. Tucker 7614 (WIS). Florida: St, George Isl., W. Calkin 146 (WIS); Sanford, Jul. 

1907/Oct. 1911 (two collections), S. Rapp s.n. (WIS). Brazil: Parana, Kalb 33324 (hb. Kalb); Mato Grosso, Kalb 

33348 (hb. Kalb). 

2. 9. 11. Thelotrema Ach., Meth. Lich.: 130 (1803). Type species: Thelotrema lepadinum 

(Ach.) Ach. 

Antrocarpum G. Mey., Nebenstunden: 326 (1825), nom. illegit. [based on Lichen lepadinus Ach.] 

Brassia Massal., Atti Reale Ist. Veneto Sci. Lett. Arti, ser. 3, 5: 259 (1860). Type species: Thelotrema 

porinoides Mont. & Bosch. 

Thelotrematomyces Thomas, in Ciferri & Tomaselli, Istit. Bot. Univ. Lab. Crittog. Pavia Atti, ser. 5, 10(1): 

52, 77 (1953), nom. illegit. 

THALLUS – Crustose, corticolous or sometimes saxicolous. Predominantly very thin to 

moderately thick, rarely distinctly thick, hyposubstratic to episubstratic, up to c. 50-800 µm 

high, grayish to greenish or olive with yellow, brown or white tones. Surface dull to slightly 

shiny, rarely shiny or ceraceous, smooth to rough or rarely pruinose, in predominantly 

hypophloedal thalli often with protuberant substrate structures, continuous to ±distinctly 

verrucose or verruculose, unfissured to fissured to rarely areolate. Prothallus thin to indistinct, 

brown. Thallus either without cortical structures or covered by a continuous to discontinuous 

protocortex up to c. 10-30 µm thick, or true cortex present, continuous to discontinuous,


hyaline to yellowish, up to c. 10-50 µm, rarely up to c. 80-100 µm thick and consisting of 

irregular to periclinal hyphae. Algal layer continuous to discontinuous, poorly to well 

developed, calcium oxalate crystals sparse to abundant, small to large, scattered to clustered, 

rarely in columnar arrangement. Distinct medulla layer absent to rarely present. Vegetative 

propagules absent in Australian species, but isidia known from the south-east Asian T. 

isidiophorum. 

ASCOMATA – Conspicuous to inconspicuous, small to very large, up to c. 0.3-2 mm in 

diam., predominantly roundish to rarely slightly irregular, only in fused ascomata appearing 

distinctly irregular, rarely slightly elongated. Peri- to apothecioid, rarely becoming 

±indistinctly chroodiscoid with age, mostly sessile to rarely erumpent, solitary to fused, nonregenerating 

to rarely regenerating, distinctly immersed to distinctly emergent, then 

hemispherical, conical, cylindrical, urceolate or subglobose, rarely annular, with same surface 

as thallus or sometimes more distinctly verruculose or verrucose. Disc not visible from 

surface to partly or rarely entirely visible, grayish to flesh-colored, rarely with white or brown 

tones, epruinose to strongly pruinose. Pores predominantly tiny to wide, up to c. 50-800 µm 

in diam., more rarely gaping, up to c. 1-1.5 mm in diam., roundish to ±irregular, entire to 

±split, proper exciple rarely not visible from surface, pore margin then formed by thalline rim, 

to predominantly becoming visible from surface, free to rarely fused, off-white to whitish in 

upper parts to sometimes yellowish or brownish towards the base, often jagged and/or 

shrunken, mostly incurved to slightly erect, rarely distinctly erect to ±recurved. Thalline rim 

margin small to gaping, thin to thick, entire to distinctly split and/or lacerate, sometimes 

eroded and/or pruinose, unlayered to ±distinctly layered, concolorous with thallus to ±brighter 

or whitish, off-white or pale brownish, incurved to ±recurved. Proper exciple predominantly 

±free, rarely fused, thin to thick, hyaline to pale yellowish internally, yellowish, orange, 

grayish or brownish marginally, sometimes with substrate or crystal inclusions, apically often 

dark-brown to rarely slightly carbonized and often covered by grayish granules, non-amyloid 

to ±distinctly amyloid at the base, rarely faintly amyloid also towards the upper parts. 

Subhymenium indistinct, evanescent to moderately thin, hyaline to yellowish or brownish. 

Hymenium non-amyloid, discoid, sometimes with a ±broader base or ±cupular, in 

perithecioid ascomata also subglobose, up to c. 80-400 µm high, predominantly non-inspersed 

and clear, sometimes inspersed, usually moderately to strongly, rarely weakly conglutinated. 

Paraphyses not thickened, rarely distinctly thin, rarely straight to usually ±bent, rarely 

distinctly curly in apical parts, parallel to ±distinctly interwoven, unbranched to rarely slightly 

branched, tips slightly to distinctly thickened, rarely not thickened, sometimes ±irregular. 

Lateral paraphyses present, conspicuous to inconspicuous, sometimes distinctly 

unconglutinated, up to c. 10-40 µm long. True columella absent, in fused ascomata rarely 

columella-like structures present. Epihymenium indistinct to ±thick, hyaline to sometimes 

grayish or brownish, rarely yellowish, egranulose to granulose. 

Asci 1-8-spored, non-amyloid, clavate, ascus walls predominantly not thickened and with a 

distinct tholus (at least in younger stages), sometimes ascus walls distinctly thickened and 

with indistinct or absent tholus. Ascospores uni- to triseriate, small to large, 10-350(400) x 5- 

50 µm, transversely septate to eumuriform. Cell walls thin to thick, smooth to sometimes 

crenate, endospore in muriform ascospores thin to thick, non-halonate to ±distinctly halonate, 

halo predominantly present in younger stages, thin to thick, sometimes ±distinctly irregular, 

hyaline to yellowish or brown, sometimes distinctly pigmented only in late stages of 

development, non- to distinctly amyloid; oblong to ellipsoid or clavi- to fusiform or 

cylindrical, rarely subglobose; with roundish to subacute, rarely distinctly acute and strongly 

tapered ends; with 4-46 x 0-10 or multiple loci, loci roundish to ±angular, subglobose, 

oblong, lentiform or ±irregular, with same shaped, hemispherical or conical end cells, 

transverse septae thin to thick, distinct to sometimes indistinct with age in densely muriform 

ascospores, regular to irregular.


PYCNIDIA – Present in several species, immersed or in thallus warts with dark pore area, 

conidia ellipsoid to oblong to rarely irregular, up to c. 2-4 x 1 µm. 

CHEMISTRY – β-Orcinol depsidones present or absent. 

ECOLOGY AND DISTRIBUTION – The Thelotrema species in Australia predominantly occur 

on tree bark and rarely on siliceous rock in altitudes ranging between sea level and 1500 m. 

The majority of species is found in rainforests, coastal forests and mangroves, rarely wet 

sclerophyll forests and shrubs, in tropical to sub-tropical climates of north-western Northern 

Territory, along the eastern coast of Queensland and northern New South Wales and on Lord 

Howe Island. Some species ±extend into warm- to cool-temperate zones from Pacific northcentral 

New South Wales to southern Victoria and Tasmania. T. lepadinum also occurs in dry 

sclerophyll forests, heath shrubs and swamplands and in Karri-Forests in Western Australia. 

At present state of knowledge, amongst the 38 species known in Australia, eleven are 

endemic (T. crespoae, T. capetribulense, T. crassisporum, T. cyphelloides, T. eungellaense, T. 

gallowayanum, T. oleosum, T. pseudosubtile, T. subadjectum, T. thesaurum, T. triseptatum), 

two are Australasian (T. circumscriptum, T. monosporum), eight are paleotropical to 

paleotemperate (T. bicavatum, T. conveniens, T. cupulare, T. foveolore, T. nostalgicum, T. 

nureliyum, T. polythecium, T. rugatulum), 15 are pansubtropical to pantropical (T. adjectum, 

T. alboolivaceum, T. defossum, T. diplotrema, T. lacteum, T. lepadodes, T. leucophthalmum, 

T. myriocarpum, T. pachysporum, T. porinaceum, T. porinoides, T. saxatile, T. saxicola, T. 

subtile, T. suecicum) and one subcosmopolitan (T. lepadinum). 

NOTES – Thelotrema is the oldest genus in the family (Acharius, 1803), with T. lepadinum 

as the first described and best known species (Acharius, 1798, as Lichen lepadinus). After 

some earlier modifications, it was more recently widely accepted to accommodate species 

with uncarbonized ascomata with lateral paraphyses (e.g., Salisbury, 1972a, 1972b [as 

Thelotrema sect. Thelotrema]; Hale, 1980, 1981). Since then, several species were excluded 

and placed in other genera: Topeliopsis (Kantvilas & Vezda, 2000), Reimnitzia (Kalb, 2001) 

and Chapsa (Frisch, 2006). Hence, in his revision, Frisch (2006) only accepted species of the 

T. lepadinum-group (=Thelotrema s. str.) in the genus, which is also accepted here in large 

parts and which is well supported by molecular data (Frisch & al., 2006; see part 3.). 

However, extended molecular analyses (see part 3.) also showed that Thelotrema is 

polyphyletic. Morphologically differing are the species ‘Thelotrema’ glaucopallens (with 

myriotremoid ascomata, placed in a separate group of uncertain taxonomic position [Frisch, 

2006]) and ‘Thelotrema’ zebrinum (with distinctly carbonized exciple, see under 

‘Leptotrema’ schizoloma-group). Not surprisingly, these taxa cluster in clades distinct from 

Thelotrema s. str. However, two species that morphologically agree well with Thelotrema s. 

str., T. bicinctulum and T. rugatulum do not cluster within Thelotrema s. str., and their 

phylogenetic placement requires further studies. Thelotrema polythecium is tentatively 

included in the genus, since it shows similarities with Chapsa, Topeliopsis as well as the O. 

clandestina-group (see also under this species). 

Thelotrema species are characterized by peri- to apothecioid ascomata with a ±free proper 

exciple and lateral paraphyses. Similar genera are Chapsa and Topeliopsis, for differences see 

there.



Thelotrema adjectum Nyl. 

Flora 49: 290 (1866). Type: Cuba, Wright ser. 2, 82 (H-Nyl. 22572-lectotype, selected by Hale [1978: 39]; 

UPS-, US!-isolectotypes). 


Fig. 102. Thelotrema adjectum: growth habit (A), ascomata (B), younger ascospores (C, D) and 

ascospore detail (E). A., B.: US-isolectotype; C.: Mangold 38 f; D.: Lumbsch & Mangold 19127 I; 

E.: Lumbsch & Mangold 19165 d. Bar= A: 2 mm; B: 0.8 mm; C, E: 10 µm; D: 12 µm. 

Thallus epi- to predominantly hypophloedal, very thin to thin, up to c. 70 µm high, 

grayish-green to pale olive. Surface dull to slightly shiny, smooth to somewhat roughened, 

continuous to very rarely slightly verrucose, fissured to unfissured. Cortex structures absent 

or covered by a predominantly thin, incontinuous protocortex up to c. 10(25) µm thick. Algal 

layer well developed, continuous, calcium oxalate very abundant, small to large, often 

clustered, incorporated in hypophloedal thallus parts. Vegetative propagules not seen. 

Ascomata inconspicuous, (moderately) small, up to c. 400 µm in diam., roundish, 

apothecioid, solitary to rarely marginally fused, immersed to slightly emergent, then 

(flattened-)hemispherical. Disc sometimes partly visible from surface, pale grayish to pale 

flesh-colored, epruinose. Proper exciple not visible from surface to apically visible, off-white, 

incurved (inner thalline rim layers might be confused with proper exciple!), pores 

(moderately) small, up to c. 200 µm in diam., roundish to irregular, entire to more often split.


Thalline rim margin moderately thick, roundish to irregular, entire to more often split to 

somewhat eroded, slightly layered, inner parts/layers off-white to pale bright-brown, outer 

parts/layers concolorous with thallus to pale (reddish-)brown due to protuberant substrate, 

incurved to erect. Proper exciple fused to apically free, moderately thin to moderately thick, 

hyaline internally, pale yellowish to pale orange marginally, sometimes with substrate 

particles incorporated, often amyloid at the base or in the lower parts. Hymenium up to c. 200 

µm high, non-inspersed, moderately conglutinated, paraphyses parallel to slightly interwoven, 

unbranched, tips unthickened to slightly thickened, lateral paraphyses present, conspicuous to 

inconspicuous, up to c. 20 µm long, columellar structures absent. Epihymenium indistinct to 

thin, hyaline, without granules or crystals, or with fine grayish granules. Asci 4-8-spored, 

tholus (moderately) thick, thin when mature. Ascospores variable, moderately small to 

moderately large, eumuriform, cell walls very thick to moderately thick with age, endospore 

(moderately) thin, often covered by a thin to rarely moderately thick, sometimes irregular 

halo, hyaline, non-amyloid to faintly amyloid, predominantly fusiform to rarely somewhat 

claviform, with predominantly narrowed-roundish ends, loci roundish to slightly angular, 

especially in younger ascospores, subglobular to oblong to somewhat irregular, transverse 

septae (moderately) thin, regular to slightly irregular, 40-80 x 8-25 µm with 12-18 x 1-6 loci. 


CHEMISTRY – Thallus K-, C-, PD-; no secondary substances detectable by TLC. 

ECOLOGY AND DISTRIBUTION – Thelotrema 

adjectum was collected in Australia on tree bark in 

tropical rainforests in altitudes ranging from sea 

level to 900 m. It is a rare species restricted to 

northern Queensland. This is the first report for 

Australia and the Paleotropics, it was previously 

known from the Neotropics, hence being a 

pantropical species. 


ecorticate thallus, the immersed to slightly 

emergent, small ascomata, the moderately large, 

muriform, hyaline, non- to weakly amyloid 

ascospores with thick to very thick cell walls and 

the absence of secondary compounds. The 

ascospore morphology is similar to T. lepadinum, 

which, however, differs in ascoma morphology, 

being emergent with entirely free proper exciple. 

For differences to T. subadjectum see under this 

species. Two other similar species lacking 


T. adjectum. 

secondary metabolites and having muriform, hyaline, large ascospores are known, viz. T. 

laceratula and T. defectum from southern U.S.A. Topeliopsis laceratula can be readily 

distinguished by the thick, yellowish true cortex and monosporic asci with larger (up to 170 

µm long), strongly amyloid ascospores. Thelotrema defectum differs by having non-layered 

ascomata and smaller (up to 40 µm long) ascospores in 2-4 per asci. 

SPECIMENS EXAMINED – Australia, Queensland: 4.5 km on Buchanan Rd., Cape Tribulation, N of Mossman, 

Hale 831361, 831386 (US). Cape Tribulation Area: Cape Tribulation NP., about half way along the Cape 

Kimberly Rd., Hale 831090, 831319 (US); Cape Kimberley, Lumbsch & Mangold 19164 h, 19165 d (F). 

Daintree NP., Mossman Gorge section, near eastern border of the park, Mangold 36 i, r (F). Atherton 

Tablelands: 14.5 km on Mulgrave River Forestry Rd., SW of Gordonvale, Hale 831652 (US); Lake Euramoo,


Lumbsch & Mangold 19127 i, Mangold 38 d, f (F); Along west boundary of Lake Eacham NP., on rd. from 

Atherton, Hale 832222 (US); Danbulla Forest Drive, E of Tinaroo Dam, 1 km W of Cathedral Fig Tree nr. 

Yungaburra, Hale 831345, 831383 (US). 2 km N of Murray Falls, W of Kennedy, Hale 831430, 831651 (US). 

Thelotrema alboolivaceum Vain. 

Mycologia 21: 38 (1929). Ocellularia alboolivacea (Vain.) Zahlbr., Catal. Lich. Univers. 8: 244 (1932). 

Type: Porto Rico, Vega Baja, Fink 2153 (TUR-Vain.34714!-holotype; FH- [invalidly lectotypified by Nagarkar 

& al. 1988], NY!-isotypes). 


Fig. 104. Thelotrema alboolivaceum: growth habit (A), ascomata (B), ascoma section (C), ascospore 

(D) and ascospores showing amyloid reaction (E). A., E.: TUR-holotype; B.-D.: Lumbsch & 

Mangold 19129 s. Bar= A: 1 mm; B: 0.3 mm; C: 200 µm; D: 13 µm; E: 10 µm. 

Thallus epi- to predominantly hypophloedal, very thin to thin, epiphloedal parts in 

ascomata area up to c. 100 µm high, olive to brownish-olive or pale yellowish-brown. Thallus 

±shiny, smooth, continuous to slightly verruculose, unfissured. Covered by ±continuous true 

cortex, consisting of periclinal to irregular hyphae, sometimes with crystal inclusions, up to c. 

35 µm thick. Algal layer poorly developed and incontinuous, calcium oxalate crystals 

abundant, small to large, often clustered. Vegetative propagules not seen. Ascomata 

conspicuous, moderately large, up to c. 700 µm in diam., roundish, apothecioid, solitary to 

marginally fused, erumpent, usually distinctly emergent, cylindrical to cone-shaped with 

flattened apex. Disc sometimes becoming partly visible from surface, pale grayish, distinctly 

pruinose. Pores small to wide, up to c. 400 µm in diam., roundish to roundish-irregular, entire 

to slightly split, proper exciple usually entirely visible from surface, free, apically whitish to 

off-white, pale brownish towards the base, incurved, often shrunken. Thalline rim margin


entire to more often and split to lacerate or eroded, sometimes ±coarsely lobed, unlayered, 

roundish, whitish to off-white, often slightly pruinose, thalline rim concolorous with thallus, 

surface often slightly verruculose, becoming erect in early stages to slightly recurved with 

age. Proper exciple free, thin, hyaline internally to yellowish-brown marginally, apically 

covered with grayish granules, non-amyloid. Hymenium up to c. 100 µm high, non-inspersed, 

moderately conglutinated, paraphyses slightly interwoven, unbranched, tips thickened, lateral 

paraphyses present, usually conspicuous, up to c. 25 µm long, columellar structures absent. 

Epihymenium thick, hyaline, with grayish granules. Asci 8-spored, tholus (moderately) thick, 

thin when mature. Ascospores small, transversely septate, very rarely with a single 

longitudinal septum, cell walls thick, sometimes slightly crenate, with moderately thick to 

moderately thin halo in younger stages, non-halonate at maturity, hyaline, distinctly amyloid, 

fusi- to claviform, ends narrowed-roundish to subacute, loci roundish to rarely slightly 

angular, subglobose to lentiform or roundish-rectangular to ±irregular, end cells 

hemispherical to more often and ±distinctly conical, septae (moderately) thick, ±regular, 15- 

23 x 5-7 µm with 4-6 (x 2) loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellowish to brown, C-, PD+ orange; containing constictic and 

stictic (majors), α-acetylconstictic and cryptostictic (traces) acids. 

ECOLOGY AND D ISTRIBUTION – Thelotrema 

alboolivaceum was collected in Australia on tree 

bark in a tropical rainforest at 700 m altitude. It is 

rare in northern Queensland. This is the first report 

for Australia. This pantropical species has been 

recorded from Central America, India (Patwardhan 

& Kulkarni, 1977) and Sri Lanka (Nagarkar, 1988). 


mostly hypophloedal, corticate thallus, the 

conspicuous, emergent ascomata with free proper 

exciple, small, transversely septate, amyloid 

ascospores with conical end cells, thickened cell 

walls and septae, and the presence of the stictic acid 

chemosyndrome. Similar taxa with transversely 

septate ascospores and stictic acid include 

T. capetribulense, T. bicinctulum, T. porinoides and 

T. triseptatum. All but T. triseptatum can be readily 

distinguished by larger ascospores (up to 50 µm 

with up to 14 loci in T. capetribulense, up to 35 µm 


T. alboolivaceum. 

with up to 11 loci in T. bicinctulum, up to 140 µm with up to 30 loci in T. porinoides), for 

differences to T. triseptatum see under this species. Two morphologically similar, stictic acid 

containing species are T. cupulare and T. leucophthalmum, which are readily distinguished by 

muriform, non-amyloid ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Atherton Tablelands, Malanda Falls, Lumbsch & Mangold 

19129 s (F). India, Karnataka, Hale 48002 (US).


Thelotrema bicavatum Nyl. 

Flora 47: 269 (1864). Ocellularia bicavata (Nyl.) Müll. Arg., Bullet Herbier Boissier 2(1): 74 (1894). Type: 

Australia ('Nova Hollandia'), with 'Lecanora subfusca var. chlarona', com. Hochstetter s.n. (H-Nyl. 22790holotype, 

M!-isotype). 


Fig. 106. Thelotrema bicavatum: growth habit (A) and ascospores (B, C). A., C.: M-isotype; B.: Hale 

830650. Bar= A: 1 mm; B: 17 µm; C: 7 µm. 

Thallus hypo- to epiphloedal, (moderately) thin, up to c. 250 µm high, pale grayish-green 

to pale yellowish-gray. Thallus surface dull, smooth, continuous to more rarely verrucose, 

slightly to distinctly fissured. Thallus covered by an incontinuous protocortex up to 20 µm 

thick. Algal layer poorly developed, incontinuous, calcium oxalate crystals sparse and small, 

scattered or in large clusters. Vegetative propagules not seen. Ascomata inconspicuous, 

moderately small, up to c. 600 µm in diam., roundish to somewhat irregular, apothecioid, 

solitary to rarely marginally fused, immersed to emergent, then hemispherical to conical to 

more rarely verrucose-urceolate. Disc partly visible from surface in mature ascomata, grayish, 

distinctly pruinose. Pores moderately small to more rarely wide, up to c. 300(500) µm in 

diam., roundish to roundish-irregular to irregular, entire to slightly split, apical to upper 

proper exciple visible from surface, free, off-white to pale yellowish, pale brownish towards 

the base, predominantly incurved to rarely somewhat erect, often ±distinctly shrunken. 

Thalline rim margin moderately thick to thin, roundish to slightly irregular, entire to slightly 

split, concolorous with thallus, thalline rim incurved to slightly erect, concolorous with thallus 

to rarely slightly brownish and with same surface. Proper exciple free, moderately thick, 

hyaline to pale yellowish internally, grayish-brown to yellowish-gray marginally, apically 

sometimes dark brownish, often with substrate particles incorporated, ±distinctly amyloid at 

the base. Hymenium up to c. 150 µm high, non-inspersed, moderately conglutinated, 

paraphyses ±straight, parallel to slightly interwoven, unbranched, tips slightly thickened, 

lateral paraphyses present, inconspicuous, up to c. 15 µm long, columellar structures absent. 

Epihymenium moderately thick, hyaline, with grayish to brownish granules. Asci 8-spored, 

tholus thick, thin when mature. Ascospores (moderately) small, transversely septate, cell 

walls and endospore (moderately) thick, hyaline, with thin halo in younger stages, faintly to 

moderately amyloid, oblong to predominantly fusi- or claviform, with rounded to subacute 

ends, loci roundish to slightly angular, oblong to lentiform, with hemispherical to conical end 

cells, septae moderately thick, ±regular, 20-40(45) x 6-9 µm with 8-14(16) loci. Pycnidia not 

seen.


CHEMISTRY – Thallus K+ orange-red, C-, PD-; containing norstictic (major) and 

connorstictic (minor to trace) acids. 


bicavatum occurs on tree bark in (sub)tropical, 

warm- and cool-temperate rainforests in altitudes 

ranging from sea level to 800 m. Although it is rare 

its is a wide-spread species occurring in northern 

and southern Queensland, north-central and central 

New South Wales and in Tasmania. This paleosubtropical 

species was previously reported from 

New Zealand (Nylander, 1888) and Japan 

(Asahina, 1931; Yasuda, 1935). 


ecorticate thallus, moderately small, immersed to 

emergent ascomata with free proper exciple, 

moderately small, transversely septate, hyaline, 

amyloid, thick-walled ascospores and the presence 

of the norstictic acid chemosyndrome. There are 

several similar taxa in Australia, all distinguished 

by the absence of lichen substances, including 

T. pseudosubtile, T. subtile and T. suecicum with 

similar ascospore size. Thelotrema suecicum further differs in the distinctly thick-walled 

immature ascospores. Thelotrema pseudosubtile has somewhat larger ascospores, up to 60 µm 

long. Thelotrema subtile has slightly larger ascospores, up to 50 µm long. The latter taxon 

was considered conspecific with T. bicavatum (Salisbury, 1972a; Matsumoto, 2000) but 

recently regarded as a distinct species (Mangold & al., 2007) predominantly due the chemical 

differences. Frisch (2006) lists T. bicavatum tentatively under T. diplotrema, see under this 

species for further differences. The identity of the T. bicavatum specimens from New Zealand 

and Japan remains uncertain, since no information on secondary metabolites of these 

collections was given. Thelotrema patwardhanii from India is the only other currently known 

Thelotrema with norstictic acid and transversely septate ascospores. It is readily distinguished 

by perithecioid ascomata and larger ascospores (up to 200 µm long). 

SPECIMENS EXAMINED – Australia, Queensland: Cardwell, Scarlett & Bell 1000 (BRI). Culpa logging area, 

SE of Tully Falls, Hale 830650 (US). Wooroi State Forest Park, W of Teewantin, Hale 830885 (US). New South 

Wales: Bulahdelah District, Myall River SF., K. & A. Kalb 18052 (hb. Kalb). Royal NP., S of Sydney, K. & A. 

Kalb 21691 (hb. Kalb). Tasmania: Maria Island NP., 1.5 km NW of summit of Mt.Maria, uppermost Counsel 

Creek, Tibell 11348 (UPS). 

Thelotrema bicinctulum Nyl. 


T. bicavatum. 

Ann. Sci. Nat. Bot., ser. 4, 15: 46 (1861b). Type: New Caledonia, Pancher s.n. (H-Nyl. 22804a!-lectotype, 

selected by Frisch [2006: 282]). 

Thelotrema terebrans Nyl., Bull. Soc. Linn. Normandie 2(7): 166 (1873). Ocellularia terebrans (Nyl.) 

Zahlbr., Cat. Lich. Univers. 2: 602 (1923). Myriotrema terebrans (Nyl.) Hale, Mycotaxon 11: 138 (1980). Type; 

India, Andaman Islands, Kurz 80 (H-Nyl. 22771!-lectotype, selected by Hale [1981: 294]; BM-, UPS!-, ZTisolectotypes). 

Ocellularia demersa Müll.Arg., Mém. Soc. Phys. Hist. Nat. Geneve 29(8): 9 (1887), nom. nov. pro Pyrenula 

clandestina Fée, Ess. Crypt. Suppl. 83 (1837), nom. illeg. [non P. clandestina Ach., 1814: 8; nec. T. 

clandestinum Fée, 1837: 6]. Thelotrema albidulum Nyl., Mém. Soc. Sci. Nat. Cherbourg 5: 118 (1857), nom.


nov. inval. [non T. albidulum Nyl., 1861: 46]. Thelotrema demersum (Müll.Arg.) Salisb., Nova Hedwigia 29: 

408 (1978). Type: 'Regio tropica - habitat non raro supra cortices Crotonis Cascarillae, L.' (G-Fée 207!lectotype, 

selected by Salisbury [1978: 408]; H-Nyl. 22768-, 22769-isolectotypes). 

Ascidium octolocularis C. Knight in Bailey, Proc. Roy. Soc. Queensland 1: 152 (1884). Ocellularia 

octolocularis (C. Knight) Shirley, Proc. Roy. Soc. Queensland 6: 188 (1889). Porina octolocularis (C. Knight) 

C. Knight in Shirley, Proc. Roy. Soc. Queensland 6: 183 (1889). Type: Australia, Queensland, Brisbane, on 

shrub trees, Bailey 204 (WELT!-lectotype, here selected; BRI-'Shirley Book', p. 22, n. 10 [BRI-AQ721230]!-, 

BRI-'Bailey Book', p. 22 [BRI-AQ720161]!-isolectotypes). 


Fig. 108. Thelotrema bicinctulum: growth habit (A, C, D), ascomata (B), ascospores (E, G) and 

ascoma and thallus section (F). A.: UPS-isolectotype of T. terebrans; B.: Lumbsch & Mangold 

19162 e; C.: A. & K. Kalb 26569: D.: BRI-isolectotype of A. octolocularis; E., F.: H-lectotype: G.: 

G-lectotype of O. demersa. Bar= A: 1.5 mm; B: 1 mm; C: 1.5 mm; D: 1 mm; E: 7 µm; F: 150 µm; 

G: 10 µm. 

Thallus variable, epi- to hypophloedal, thin to more rarely thick, c. up to 500(800) µm 

high, pale grayish-green to pale yellowish-gray or (pale) olive. Surface dull to shiny, smooth 

to sometimes partly porous-rough, continuous to verrucose, unfissured to strongly fissured or


rimose, then upper thallus usually partly flaking away, exposing the whitish medulla. True 

cortex present, very variably developed, continuous to sometimes incontinuous or lacking in 

large parts, consisting of irregular to periclinal hyphae, up to c. 35 µm thick. Algal layer 

mostly ±well developed, continuous, sometimes becoming incontinuous due to large crystal 

inclusions, calcium oxalate crystals very abundant, often almost filling the entire lower 

thallus, variable in size, in clusters or scattered, distinct whitish medulla layer present. 

Vegetative propagules not seen. Ascomata ±inconspicuous, moderately small, up to c. 400 

µm in diam., roundish, apothecioid, predominantly solitary, rarely slightly fused, 

predominantly ±immersed. Disc often becoming partly visible, grayish, coarsely pruinose. 

Pores small, up to c. 120 µm in diam., roundish to somewhat irregular, apical to upper proper 

exciple visible from surface, free, entire to slightly split, sometimes ±jagged, off-white to 

whitish, predominantly incurved to rarely somewhat erect, sometimes ±distinctly shrunken. 

Thalline rim margin (moderately) thin, roundish to slightly irregular, entire to slightly split, 

concolorous with thallus to brighter than thallus, often somewhat raised, thalline rim 

incurved, concolorous with thallus. Proper exciple free in the upper parts, (moderately) thin, 

hyaline internally to pale yellowish to dark yellowish-brown marginally, apically often 

covered by grayish granules, non-amyloid to sometimes amyloid at the base. Hymenium up to 

c. 100 µm high, non-inspersed, moderately conglutinated, paraphyses straight to slightly bent, 

slightly interwoven, unbranched, tips slightly thickened, lateral paraphyses present, often 

inconspicuous, up to c. 25 µm long, columellar structures absent. Epihymenium usually thick, 

hyaline, with grayish granules. Asci 8-spored, tholus moderately thick, thinning or not visible 

at maturity. Ascospores variable, small, transversely septate, cell walls thin to moderately 

thick, usually with thin halo, hyaline, non-amyloid to faintly amyloid, narrow, ellipsoid to 

fusi- or claviform with roundish to subacute ends, loci roundish to angular, predominantly 

oblong to lentiform or rectangular, with hemispherical to conical end cells, septae moderately 

thin, regular, 20-35 x 5-7 µm with 8-11 loci. Pycnidia not seen. 


constictic, hypostictic (minor to trace), α-acetylconstictic, hypoconstictic, cryptostictic and αacetylhypoconstictic 



bicinctulum was collected in Australia on tree bark 

in (sub)tropical rainforests, mangroves, coastal 

forests and wet sclerophyll forests in altitudes 

ranging from sea level to 800 m. It is a moderately 

common and wide-spread species occurring in 

north-western Northern Territory and Queensland. 

Besides Australia it was reported from Africa 

(Frisch, 2006), India (Hale, 1981), Sri Lanka 

(ibid.), India (ibid.), Andman Islands (Nagarkar & 

al., 1988) and New Caledonia indicating a 

paleo(sub)tropical distribution. 

N OTES – It is a variable species that is 

characterized by the ±corticate thallus with 

abundant crystals, the small ascomata with free 

proper exciple and ±inconspicuous lateral 

paraphyses, moderately small, transversely septate, 

hyaline, narrow, non-amyloid to faintly amyloid 

ascospores and the stictic acid chemosyndrome. 


T. bicinctulum.


Similar are T. alboolivaceum, T. capetribulense and T. triseptatum, for differences see under 

these species. Thelotrema euphorbia from Africa is also similar but no type material was 

available for study (Frisch, 2006). Two collections are only tentatively identified as T. 

bicinctulum here: Kalb 21620 has distinctly larger ascospores, up to 45 µm long with up to 14 

loci, and Hale 832714 has distinctly submuriform ascospores. Four Australian collections are 

sterile and hence only tentatively identified as well: Hale 832151, 832434, Mangold 38 a, 

Wilson 1450. 

SPECIMENS EXAMINED – Australia, Northern Territory: Doctors Gully, 2 km NW of Darwin, Streimann 8762 

(CANB). Queensland: Near Cedar Bay NP., on rd. to Cooktown, Mangold 34 f (F). NW of Mossman, Km 45 on 

Mt. Windsor Rd., Hale 832458 (US). Cape Tribulation Area: Myall Beach, Lumbsch & Mangold 19161 q, 

19162 c, e (F); Track to Cape Tribulation Beach, Mangold 32 a, c (F). Kuranda Range, SE of Kuranda, K. & A. 

Kalb 21620 (hb. Kalb). W of Palm Cove, 25 km N of Cairns, K. & A. Kalb 19972 (hb. Kalb). Manchans Beech, 

N of Cairns, K. & A. Kalb 21182 (hb. Kalb). Atherton Tablelands, Lake Euramoo, Mangold 38 a (F). 13 km 

from Koombooloomba rd. turnoff, SE of Tully Falls, Hale 832434 (US). Dawson logging area, SF. Reserve 605, 

24 km S of Koombooloomba turnoff, WSW of Tully, Hale 832714 (US). Conway Range NP., near Shute 

Harbour-Airlie Beach, Hale 832421 (US). Cape Hillsborough NP., NW of Mackay: Hale 832513 (US); Along 

the walking track from the Casuarina Beach to Beachcombers cove, Thor 5138 (S). Frasier Coast, River Heads, 

Lumbsch & Mangold 19092 c, j, l, n (F). Noosa NP., Palm Grove Track, Hale 831707 (US). Mt. Mee SF., NW 

of Mt. Mee, Hale 832151 (US). Mt. Glorious, Brisbane SF., 40 km WNW of Brisbane, K. & A. Kalb 26569 (hb. 

Kalb). Goodna (near Brisbane), Wilson 1450, pr. p. (NSW). 

Thelotrema capetribulense Mangold spec. nov. ined. 

Type: Australia, Queensland: Cape Tribulation Area, Myall Beach, Lumbsch & Mangold 19161 x (CANBholotype; 

BRI-, F-isotypes). 

ETYMOLOGY – The epithet refers to Cape Tribulation in northern Queensland where the 

species is most common. 


Thallus epi- to hypophloedal, thin to moderately thick, up to c. 400 µm high, grayish to 

pale grayish-green. Surface slightly shiny, smooth, distinctly verrucose, unfissured to slightly 

fissured. Cortex structures variable, predominantly an ±incontinuous protocortex present, up 

to c. 30µm thick, in some parts becoming distinctly conglutinated forming a true cortex of 

irregular to periclinal hyphae. Algal layer poorly developed, usually incontinuous due to 

calcium oxalate crystal inclusions, calcium oxalate crystals abundant, small to more often 

large, scattered or clustered. Vegetative propagules not seen. Ascomata conspicuous, 

moderately large, up to c. 700 µm in diam., roundish, apothecioid, solitary to sometimes 

slightly fused marginally, usually distinctly emergent, predominantly subglobose to urceolate 

to more rarely broad-cylindrical with ±verrucose surface. Disc not visible from surface to 

rarely becoming somewhat visible, pale brownish-gray, indistinctly pruinose. Pores small to 

more rarely moderately wide, up to c. 300 µm in diam., roundish, entire to split, proper 

exciple entirely to apically visible from surface, becoming free, apically bright, darker 

towards the base, predominantly incurved. Thalline rim margin thick, entire to somewhat split 

to slightly lacerate or eroded, often slightly layered, ±roundish, mostly whitish or brighter 

than thallus, sometimes slightly pruinose, incurved to more rarely erect. Proper exciple 

becoming free, ±thin, hyaline internally to pale yellowish-brown marginally, amyloid at the 

base. Hymenium up to c. 150 µm high, non-inspersed, moderately conglutinated, paraphyses 

interwoven, unbranched, tips slightly thickened, lateral paraphyses present, usually 

conspicuous, up to c. 30 µm long, columellar structures absent. Epihymenium inconspicuous, 

thin, hyaline, with grayish granules. Asci 8-spored, tholus thick to moderately thin at


Fig. 110. Thelotrema capetribulense: growth habit (A) and young and mature ascospores (B, C). A.- 

C.: CANB-holotype. Bar= A: 0.5 mm; B: 7 µm; C: 4 µm. 

maturity. Ascospores typical, (moderately) small, 

transversely septate, cell walls (moderately) thick, 

with thin, sometimes irregular halo, hyaline, 

moderately amyloid, fusi- to more often claviform, 

ends narrowed-roundish to subacute, loci irregular 

and variable, roundish to moderately angular, 

subglobose to lentiform, end cells hemispherical to 

conical, septae (moderately) thick, regular to 

±irregular, 25-50 x 5-10 µm with 6-14 loci. 



C-, PD+ orange; containing constictic, stictic, 

hypostictic (majors), hypoconstictic and cryptostictic 



capetribulense occurs on tree bark in tropical 

rainforests and mangroves in altitudes ranging from 

10 to 700 m. It is rare and currently only known 

from northern Queensland. 


T. capetribulense.


NOTES – This new species is characterized by the verrucose, moderately thick thallus, the 

conspicuous, emergent ascomata with free proper exciple, medium-sized, amyloid ascospores 

with thick cell walls and septae and the presence of the stictic acid chemosyndrome. It is 

similar to T. porinoides, for differences see under this species. Morphologically similar 

ascospores are found in T. triseptatum and T. suecicum. These two species differ in having 

usually smaller ascospores. Thelotrema triseptatum also contains the stictic acid 

chemosyndrome, while T. suecicum lacks secondary metabolites. Thelotrema bicinctulum can 

be readily distinguished by smaller (up to 400 µm in diam.), immersed ascomata and smaller 

ascospores (up to 35 µm long with up to 11 loci) with thinner walls and septae. 

SPECIMENS EXAMINED – Australia, Queensland: Cape Tribulation Area, Myall Beach, Lumbsch & Mangold 

19158 u, 19160 q, y, 19162 i (F). Atherton Tablelands, Lamb Range, N of Gillies Rd., 22 km ENE of Atherton, 

Thor 5738 pr.p. (S). 

Thelotrema circumscriptum C. Knight 

Trans. New Zealand Inst. 15: 349 (1883). Ocellularia circumscripta (Knight) Dodge, Nova Hedwigia 19: 

489 (1970). Type: New Zealand, s.l. (?Wellington), 1883, C. Knight s.n. (WELT-Knight 35A: 2-lectotype, 

selected by Galloway [1985: 573]; G!-isolectotype). 


Fig. 112. Thelotrema circumscriptum: growth habit (A), ascomata (B), ascospores (C) and ascospores 

showing amyloid reaction (D). A.-D.: G-isolectotype. Bar= A: 1.5 mm; B: 0.6 mm; C: 10 µm; D: 8 

µm. 

Thallus epi- to hypophloedal, thin to moderately thick, up to c. 300 µm high, pale grayishgreen 

to pale olive. Surface dull to slightly shiny, smooth, continuous to slightly verrucose, 

slightly to distinctly fissured. Cortex structures variable, predominantly an ±continuous


protocortex present, up to c. 25 µm thick, in some parts becoming distinctly conglutinated 

forming a true cortex consisting of irregular hyphae. Algal layer well developed and 

continuous, becoming somewhat incontinuous due to large calcium oxalate crystal inclusions, 

calcium oxalate crystals moderately abundant, usually large, scattered to more rarely 

clustered. Vegetative propagules not seen. Ascomata conspicuous, moderately large, up to c. 

800 µm in diam., roundish, apothecioid, solitary to sometimes fused, becoming distinctly 

emergent, predominantly hemispherical with same surface as thallus. Disc not visible from 

surface to sometimes becoming partly visible, pale grayish, indistinctly pruinose. Pores 

(moderately) small, up to c. 200 µm in diam., roundish, entire to split, proper exciple apically 

more rarely entirely visible from surface, free, often shrunken, apically bright, darker towards 

the base, predominantly incurved. Thalline rim margin moderately thin to moderately thick, 

predominantly entire, ±roundish to more rarely slightly elongate, incurved, usually slightly 

brighter than thallus. Proper exciple becoming free, thin, hyaline internally to pale brownish 

marginally, apically often with grayish granules, often moderately amyloid at the base. 


interwoven, unbranched, tips distinctly, irregularly thickened, lateral paraphyses present, 

sometimes inconspicuous, up to c. 20 µm long, columellar structures absent. Epihymenium 

inconspicuous, thin, hyaline, with small grayish to colorless granules. Asci 8-spored, tholus 

thick, thin when mature. Ascospores typical, (moderately) small, transversely septate, cell 

walls (moderately) thick, with thin halo, hyaline, moderately to distinctly amyloid, fusi- to 

more often claviform, ends narrowed-roundish to subacute, loci variable, roundish to 

moderately angular, subglobose to lentiform, end cells hemispherical to conical, septae 

(moderately) thick, slightly irregular, 25-45 x 7-9 µm with 8-14(15) loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellow becoming red, C-, PD+ orange; containing salazinic acid. 


circumscriptum was collected in Australia on bark 

of a dead palm in a subtropical lowland rainforest 

at 170 m. In Australia it is only known from Lord 

Howe Island. This is the first report for Australia. It 

has previously been known from New Zealand, 

being an Australasian element. 


moderately thick, predominantly corticate thallus, 

emergent ascomata with free proper exciple, 

moderately small, transversely septate, hyaline, 

amyloid ascospores with thick cell walls and septae 

and salazinic acid as secondary metabolite. It is 

similar to the salazinic acid containing T. hians, 

which can be distinguished by the thinner thallus, 

the reddish-brown, more gaping thalline rim 

margin, a somewhat thicker, apically more 

distinctly darkened exciple, and larger and broader 

ascospores (up to 80 x 12 µm in size with up to 16 


T. circumscriptum. 

loci). Another similar Australian species with almost identical ascospores is the tropical T. 

capetribulense, which differs in containing the stictic acid chemosyndrome and a thicker, split 

to lacerate, sometimes layered, distinctly bright thalline rim margin. Thelotrema subtile can 

be distinguished by a usually thinner thallus, ascospores that turn brownish in over-mature or 

decayed stages, and the absence of secondary compounds. Zahlbruckner (1924) considered T.


circumscriptum as conspecific with O. cavata, a distinct taxon with carbonized exciple and a 

columella. 

SPECIMENS EXAMINED – Australia, New South Wales: Lord Howe Island, Smoking Tree Ridge, Elix 42148 

(CANB). New Zealand, nr. Wellington, Buchanan 141 pr. p. [with lectotype of T. hians] (GLAM). 

Thelotrema conveniens Nyl. 

Bull. Soc. Linn. Normand. 2(7): 168 (1873). Type: Colombia ("Nova Granata"), Tequendama, 1863, Lindig 

s.n. (H-Nyl. 22496!-lectotype, selected by Hale [1978: 42]). 


Fig. 114. Thelotrema conveniens: growth habit (A), ascomata (B), ascoma section (C), ascospores (D, 

F), over-mature ascospore (E) and ascospore showing amyloid reaction (G). A., C., E.-G.: Hlectotype; 

B., D.: Tibell 12638. Bar= A: 2 mm; B: 0.75 mm; C: 150 µm; D: 35 µm; E-G: 25 µm. 

Thallus variable, corticolous to rarely saxicolous, epi- to predominantly hypophloedal, 

predominantly epilithic in saxicolous specimen, very thin to thin to more rarely moderately 

thick, up to c. 300 µm high, in shades of gray to pale yellowish-brown or pale grayish-green. 

Surface variable, dull to slightly shiny, smooth to roughened, often due to protuberant


substrate, continuous to slightly verrucose, ±fissured to sometimes areolate. True cortex 

usually absent, thallus predominantly covered by an continuous to incontinuous protocortex 

up to 25 µm thick, rarely becoming weakly conglutinated forming a true cortex of periclinal 

to irregular hyphae. Algal layer variable, well to poorly developed, predominantly 

incontinuous, calcium oxalate usually abundant, rarely sparse, small to large, scattered or 

clustered. Vegetative propagules not seen. Ascomata variable, conspicuous to more often 

inconspicuous, moderately small to moderately large, up to 600 µm in diam., ±roundish, perito 

predominantly apothecioid, solitary to marginally slightly fused, immersed to ±distinctly 

emergent, (verrucose-)hemispherical to more rarely (verrucose-)urceolate. Disc sometimes 

becoming partly visible from surface, pale grayish to gray, distinctly pruinose. Pores variable, 

small to wide, up to c. 300 µm in diam., roundish to more often roundish-irregular to 

irregular, proper exciple rarely entirely to more often only apically visible from surface, rarely 

entirely free to more often free only in upper parts, off-white to whitish, ±shrunken, 

predominantly incurved to rarely somewhat erect. Thalline rim margin roundish to irregular, 

small to gaping, entire to ±distinctly split, rarely somewhat eroded, moderately thin to 

moderately thick, incurved to rarely slightly erect, concolorous with thallus to sometimes 

brighter than thallus or brownish. Proper exciple free usually only in the upper parts, 

moderately thin to moderately thick, hyaline to pale yellowish internally, yellowish-brown to 

sometimes brownish marginally, apically sometimes dark-brown to dark-gray, very rarely 

slightly amyloid at the base and in parts of subhymenium. Hymenium up to c. 250 µm high, 

non-inspersed, moderately conglutinated, paraphyses parallel or slightly interwoven, 

unbranched, tips moderately thickened, lateral paraphyses present, sometimes inconspicuous, 

up to c. 30 µm long, columellar structures absent. Epihymenium variable, indistinct to 

moderately thin to thick, hyaline, without granules or usually with fine to coarse grayish to 

dark-gray granules. Asci 1-spored, tholus (moderately) thick, moderately thin when mature. 

Ascospores moderately to very large, densely eumuriform, cell walls and endospore 

predominantly (moderately) thin, only in younger stages and early maturity sometimes 

distinctly thickened parts present, non-halonate, hyaline becoming yellowish to brownish with 

age, distinctly brown usually only at late stages of maturity or in decayed ascospores, 

distinctly amyloid at maturity, showing a very strong amyloid reaction at maturity (before 

becoming distinctly pigmented), cylindrical to oblong-ellipsoid or broad-fusiform to more 

rarely bifusiform, with roundish to narrowed-roundish ends, loci predominantly roundish to 

slightly angular, subglobular to ±irregular, 

transverse septae thin, distinct and regular 

throughout development, 80-200 x 20-40 µm with 

multiple loci. Pycnidia not seen. 




conveniens was collected in Australia on tree bark 

and on rock, in (sub)tropical and warm-temperate 


It is moderately common and wide-spread 

occurring in northern and southern Queensland and 

in south-central New South Wales. This is the first 

report for Australia and the paleotropics (see notes 

below). Previously this pan(sub)tropical species 

was known from the Neotropics. 


T. conveniens.


N OTES – Thelotrema conveniens is a variable taxon that is characterized by a 

predominantly thin and ecorticate thallus, immersed to emergent ascomata with apically free 

exciple, monosporic asci, large, desnely muriform ascospores and the absence of secondary 

compounds. It is similar to T. monosporum and T. saxatile. Thelotrema conveniens can be 

readily distinguished by less distinctly pigmented ascospores. Thelotrema monosporum 

differs in smaller (up to 100 µm long) ascospores in 2-4-spored asci. Another similar species 

is T. rugatulum, which differs in unpigmented ascospores and 2-spored asci. For differences 

to T. lepadodes see under that species. Parts of the Australian collections are somewhat 

different from the Columbian type, which has a thicker thallus that is covered by more 

distinct cortex structures and has more distinctly emergent ascomata with smaller pores, 

usually not exposing the disc. However, several Australian specimens show intermediate 

morphs. 

SPECIMENS EXAMINED – Australia, Queensland: Atherton Tablelands: Danbulla Forest Drive: 4 km E of 

Tinaroo, Hale 8325339 (US); Near Lake Euramoo parking lot, Mangold 37 c (F); Lake Eacham NP., Mangold 

29 af (F). 5 km N of Yarraman and Yarraman Forest Drive, Hale 831397 (US). 6 km N of Jimna, Tibell 12795, 

12638 (UPS). 25 km WNW of Nambour, 3 km N of Kenilworth, Tibell 12638 (UPS). Mt. Mee SF., 6 km NW of 

Forestry Office, NW of Mt. Mee, Hale 830617, 830838, 830839 (US). Darling Downs [Brisbane area], 1893, 

Bailey s.n. (G-10194/13, -10194/14, BRI-AQ721254 ['Shirley Book', p. 23, n. 14]). Head of Teviot Brook, NE of 

the Head, Boonah rd., Hale 59485 (US). Cunninghams Gap NP., lower slopes of Mt. Cordeaux exposed to the 

south, Hafellner 16234, 16277 (GZU). Lamington NP., Python Rock Track, Hale 832430 (US). New South 

Wales: Bairne Track, 30 km N of Sydney, 30.09.1989, Archer 'P 62' (HO). Uncertain location [?Queensland], 

(as Porina praestantior var. nana), Bailey 802 (BRI). New Caledonia, Yaté, Hill 11661 (US). 

Thelotrema crassisporum Mangold spec. nov. ined. 

Type: Australia, Queensland, Eungella National Park, Tibell 14717 (UPS-holotype). 

ETYMOLOGY – The epithet refers to the form of the ascospores (from lat. crassus =thick). 


Thallus predominantly hypophloedal, epiphloedal usually only in ascomata area, thin, up 

to c. 100 µm high, pale yellowish gray to pale greenish gray. Surface dull to slightly shiny, 

smooth, continuous to slightly verrucose, unfissered, appearing somewhat fissured due to 

substrate structure. True cortex present, continuous to somewhat incontinuous, up to 30 µm 

thick, consisting of periclinal hyphae, usually with incorporated substrate particles. Algal 

layer well developed, continuous to incontinuous, calcium oxalate crystals lacking to sparse, 

large. Vegetative propagules not seen. Ascomata conspicuous, moderately large, up to c. 600 

µm in diam., roundish, apothecioid, solitary to slightly marginally fused, emergent, 

predominantly hemispherical. Disc not visible from surface to rarely partly visible, pale 

grayish, slightly pruinose. Pores small, up to c. 100 µm in diam., irregular, proper exciple 

visible from surface, free, off-white to pale grayish, incurved and distinctly shrunken. 

Thalline rim margin moderately thin to moderately thick, entire, roundish to slightly irregular, 

thalline rim incurved, pale reddish brown, with same surface as thallus. Proper exciple usually 

entirely free, thin, hyaline internally to predominantly yellowish-brown marginally, apically 

usually dark-brown, non-amyloid. Epihymenium indistinct to thin, hyaline, with sparse 

grayish granules, without crystals. Asci 8-spored, tholus (moderately) thick, moderately thin 

when mature. Ascospores small, (sub-)muriform, cell walls moderately thin to moderately 

thick, endospore thin, non-halonate, hyaline, non-amyloid, subglobose to ellipsoid to more 

rarely oblong, with predominantly roundish ends, loci roundish to angular, (roundish-)cubical


Fig. 116. Thelotrema crassisporum: growth habit (A), ascomata (B), and ascospores (D-E). A.-E.: 

UPS-holotype. Bar= A: 2 mm; B: 0.4 mm; C: 15 

µm; D, E: 3.5 µm. 

to irregular, transverse septae (moderately) thin, in 

immature ascospores irregular, becoming ±regular 

with maturity, 12-20 x 8-12 µm with 4-7 x 1-4 loci. 



C-, PD+ orange; containing constictic and stictic 

(majors) acids. 


crassisporum was collected in Australia on a soft 

bark of an unknown tree in a tropical rainforest. It 

is only known from the type locality in northcentral 

Queensland. 


hypophloedal thallus, large, emergent, ascomata 


T. crassisporum.


with free exciple, small, hyaline, (sub-)muriform, non-amyloid ascospores and the stictic acid 

chemosyndrome. It is similar to the temperate T. lepadinum that can be distinguished by the 

lack of secondary compounds and distinctly larger ascospores (up to 140 µm long). 

Thelotrema myriocarpum is another similar species, which can be readily distinguished by the 

thicker thallus and the predominantly immersed, smaller ascomata with only partly free 

exciple and larger ascospores (up to 40 µm long). Thelotrema cyphelloides agrees in having 

stictic acid and similar ascospores but differs by a dull, roughened, ecorticate thallus and 

smaller, immersed ascomata. 

SPECIMENS EXAMINED – See type collection of this species. 

Thelotrema crespoae Mangold spec. nov. ined. 

Type: Australia, Queensland, Wooroi State Forest Park, W of Teewantin, Hale 832786 (US-holotype). 

ETYMOLOGY – This new species is dedicated to the Spanish lichenologist Ana Crespo. 

Fig. 118. Thelotrema crespoae: growth habit (A), ascomata (B), ascoma section (C), ascospore 

(D), and ascospore details (E, F). A., C.-F.: US-holotype; B.: Mangold 27 v. Bar= A: 0.8 mm; 

B: 0.6 mm; C: 200 µm; D: 40 µm; E: 12 µm; F: 20 µm.



Thallus epi- to hypophloedal, thin, up to c. 150 µm high, pale grayish-green to grayish. 

Surface dull, roughened, continuous, often with protuberant substrate structure, unfissured 

(appearing fissured due to substrate structure). Cortex structures absent. Algal layer 

moderately to poorly developed, incontinuous, calcium oxalate crystals predominantly 

abundant, usually small, scattered to more rarely clustered. Vegetative propagules not seen. 

Ascomata inconspicuous, (moderately) large, up to c. 800 µm in diam., roundish, perithecioid 

in younger stages, becoming apothecioid with age, solitary, moderately to distinctly emergent 

at maturity, hemispherical to urceolate. Disc usually not visible from surface, rarely becoming 

somewhat visible, grayish, pruinose. Pores small to moderately wide, up to c. 400 µm in 

diam., roundish to slightly irregular, entire to slightly split, apical proper exciple becoming 

visible from surface, often somewhat shrunken, off-white, pale brownish towards the base, 

incurved to slightly erect. Thalline rim margin moderately small, becoming moderately wide 

to gaping with age, moderately thin to thick, predominantly split, sometimes rugged to 

somewhat lacerate, thalline rim incurved to slightly erect, concolorous with thallus to more 

rarely brownish. Proper exciple becoming entirely free, predominantly moderately thin, 

hyaline internally to (pale)brownish or yellowish-brown marginally, apically sometimes with 

coarse grayish granules, non-amyloid. Hymenium up to c. 300 µm high, non-inspersed, 

moderately conglutinated, paraphyses parallel to somewhat interwoven, unbranched, tips 

slightly to distinctly thickened, lateral paraphyses present, inconspicuous, up to c. 20 µm 

long, columellar structures absent. Epihymenium (moderately) thick, hyaline, with (dark) 

grayish granules. Asci single-spored, tholus moderately thick, not visible at maturity. 

Ascospores typical, very large, transversely septate, cell walls thick, non-halonate, ±distinctly 

brown at late stage, non-amyloid to distinctly amyloid at early maturity (just before initial 

pigmentation), oblong-fusiform, ends characteristic, distinctly acute, appendix-like in younger 

stages to strongly tapered in older ascospores, loci roundish to somewhat acute, oblong to 

(acute-)lentiform, loci in tapered areas ±rectangular to irregular, end cells distinctly conical, 

septae thin to thickened in late maturity, distinctly regular, 150-280 x 25-35 µm with multiple 





crespoae was collected in Australia on tree bark in 

a subtropical and in a warm-temperate coastal 

rainforest in altitudes ranging from 10 to 50 m. It is 

a rare species, currently only known from southern 

Queensland and central New South Wales. 

NOTES – This new species is easily recognized 

by the large (up to 280 µm), transversely septate, 

brownish ascospores with strongly acute, usually 

appendix-like tips. The most similar species is 

T. lacteum, which is readily distinguished by 

smaller (up to 130 µm) ascospores in 4-8-spored 

asci. 

SPECIMENS EXAMINED – Australia, New South Wales: 

Myall Lakes NP., Mungo Brush Camping Area, Mangold 27 v 

(F). 


T. crespoae.


Thelotrema cupulare Müll.Arg. 

Hedwigia 32: 131 (1893). Type: Australia, Queensland, Brisbane, Bailey 375 (G!-holotype; BRI-'Shirley 

Book', p. 22, n. 20 [BRI-AQ721240]!-isotype). 

Thelotrema dissultum Hale, Bull. Br. Mus. nat. Hist. (Bot.) 8(3): 254 (1981). Type: Sri Lanka, 

Sabaragamuwa, Hale 51139 (US!-holotype, BM-isotype). 


Fig. 120. Thelotrema cupulare: growth habit (A), ascomata (B) and ascospores (C-E). A.: Hale 

831595; B.: G-holotype; C.: A. & K. Kalb 19834; D.: US-holotype of T. dissultum; E.: Hale 

831591. Bar= A: 2 mm; B: 1.2 mm; C: 4 µm; D, E: 6 µm. 

Thallus epi- to hypophloedal, thin to moderately thick, up to c. 200 µm high, yellowishbrown 

to olive. Surface waxy, smooth, continuous, unfissured, sometimes appearing fissured 

due to substrate structure. True cortex present, yellowish, up to c. 100 µm thick, consisting of 

irregular to periclinal hyphae. Algal layer continuous to incontinuous, well developed, 

calcium oxalate crystals abundant, large to more often small, scattered to clustered. 

Vegetative propagules not seen. Ascomata very conspicuous, large, up to c. 1.8 mm in diam., 

roundish, ±irregular in fused ascomata, erumpent, apothecioid to sometimes chroodiscoid, 

solitary to distinctly fused, sometimes regenerating, immersed to moderately emergent, then 

depressed-cylindrical to irregular. Disc partly to sometimes entirely visible from surface, pale 

grayish to pale flesh-colored, coarsely pruinose. Pores becoming wide to gaping, up to


c. 1 mm in diam., roundish to moderately irregular, inner and apical proper exciple becoming 

visible from surface, fused to free, ± jagged, whitish, incurved to recurved, somewhat 

shrunken in entirely free proper exciples. Thalline rim margin predominantly thick, roundish 

to moderately irregular, split to lacerate, ±coarsely lobed, sometimes indistinctly layered, 

becoming ±distinctly eroded with age, whitish, pruinose, thalline rim concolorous and with 

same surface as thallus, predominantly erect to recurved. Proper exciple partly to distinctly 

free in older stages, moderately thick, hyaline internally to (pale)yellowish-gray or brownish 

marginally, often with calcium oxalate crystal inclusions, apically usually covered by grayish 

granules, non-amyloid to rarely faintly amyloid at the base. Hymenium up to c. 100 µm high, 

non-inspersed, moderately conglutinated, paraphyses ±bent, ±interwoven, unbranched, tips 

slightly thickened and somewhat irregular, lateral paraphyses present, usually very 

inconspicuous, up to c. 20 µm long, true columella absent, in fused ascomata sometimes with 

columella-like structures. Epihymenium ±thick, hyaline to pale grayish-brown, with grayish 

granules and sometimes small crystals. Asci 8-spored, tholus (moderately) thick, thin when 

mature. Ascospores small, transversely septate when young, becoming (sub-)muriform in late 

stage, cell walls moderately thin to moderately thick, endospore moderately thick, nonhalonate, 

hyaline, non-amyloid, ellipsoid to fusi- or claviform, with roundish to subacute 

ends, loci roundish to slightly angular, subglobose to lentiform, transverse septae moderately 

thick, regular, 15-25(30) x 5-8(12) µm with 4-9(10) x 1-3(4) loci. Pycnidia not seen. 


constictic (major to minor) and α-acetylconstictic (trace) acids. 


cupulare occurs on tree bark in (sub)tropical 

rainforests in altitudes ranging from sea level to 

500 m. This rare species occurs in northern and 

southern Queensland. Previously it was recorded 

from Sri Lanka. 

NOTES – This taxon is characterized by a dark, 

waxy thallus with a true cortex, large ascomata 

with a whitish, lacerate to eroded, ±pruinose 

thalline rim margin, moderately small, (sub-) 

muriform, hyaline, non-amyloid ascospores and the 

presence of the stictic acid chemosydrome. It is 

similar to T. leucophthalmum and T. thesaurum, 

which both differ in larger ascospores (up to 60 µm 

long with up to 16 x 6 loci in T. leucophthalmum, 

up to 130 µm long in T. thesaurum). Thelotrema 

paralbidum from the Bahamas is also similar, but 

the thallus is ecorticate and the ascospores are more 

distinctly muriform and have smaller loci. 


T. cupulare. 

SPECIMENS EXAMINED – Australia, Queensland: Kuranda Range, NW of Cairns, A. & K. Kalb 19834 (hb. 

Kalb). Kalpowar Forest Drive, 40 km NE of Monto, SW of Gladstone, Hale 831591 (US). Brisbane: Bailey 278 

(BM, BRI); s.c. [?Bailey] (F).


Thelotrema cyphelloides Müll.Arg. 

Bull. Herb. Boissier 3: 314 (1895). Type: Australia, Queensland, Brisbane, Bailey 700 (G!-holotype, BRI- 

'Shirley Book', p. 22, n. 17 [BRI-AQ721237]!-isotype). 


Fig. 122. Thelotrema cyphelloides: growth habit (A), ascoma and thallus section (B) and ascospores 

(C, D). A.: G-holotype: C.-D.: BRI-isotype. Bar= A: 1 mm; B: 150 µm; C: 12.5 µm; D: 5 µm. 

Thallus predominantly hypophloedal, thin, up to c. 100 µm high, pale grayish-green. 

Surface dull, rough, continuous to verruculose, strongly fissured. Cortex structures absent. 

Algal layer well developed and continuous, calcium oxalate crystals absent to very sparse, if 

present, large. Vegetative propagules not seen. Ascomata conspicuous, moderately small, up 

to 300 µm in diam., roundish to slightly irregular, apothecioid, solitary, immersed. Disc often 

partly visible from surface, pale flesh-colored to orange, epruinose. Pores (moderately) small, 

up to c. 250 µm in diam., roundish to sometimes slightly irregular, upper proper exciple 

visible from surface, free, entire to more rarely slightly split, ±whitish, incurved to erect. 

Thalline rim margin moderately thick, roundish to slightly irregular, entire to somewhat 

eroded, concolorous with thallus, margin area sometimes circularly fissured, somewhat 

resembling a layered margin, thalline rim indistinctly incurved to erect, not distinguishable 

from main thallus. Proper exciple apically free, moderately thin, hyaline to pale yellowish 

internally, pale orange marginally, non-amyloid. Hymenium up to c. 100 µm high, noninspersed, 

distinctly conglutinated, paraphyses straight to slightly bent, slightly interwoven, 

unbranched, tips slightly thickened, lateral paraphyses present, inconspicuous, up to c. 15 µm


long, columellar structures absent. Epihymenium indistinct, hyaline, without granules or 

crystals. Asci 8-spored, tholus thick, becoming moderately thin when mature. Ascospores 

small, (sub-)muriform, cell walls moderately thick, endospore (moderately) thick, nonhalonate, 

hyaline, non-amyloid to faintly amyloid in older stages, oblong to ellipsoid to 

somewhat claviform, with roundish to narrowed-roundish ends, loci predominantly roundish, 

subglobose to oblong, transverse septae moderately thick, regular, 20-27 x 8-10 µm with 7-9 

x 1-4 loci. Pycnidia not seen. 


stictic, (majors), and hypostictic (minor) acids. 


cyphelloides was collected in Australia on resinous 

bark. It is only known from the type from southern 

Queensland, which has no further information 

concerning the habitat. 


ecorticate, dull, rough and strongly fissured thallus, 

immersed ascomata with relatively wide pores and 

a free proper exciple, small, hyaline, (sub-) 

muriform, thick-walled ascospores and the stictic 

acid chemosyndrome. It is similar to T. myriocarpum, 

but differs morphologically. Thelotrema 

myriocarpum has a shiny, smooth, corticate, nonfissured 

thallus. Another similar Australian species 

with stictic acid and small, hyaline, muriform, nonamyloid 

ascospores is T. crassisporum, which has 

large, emergent ascomata and smaller (up to 15 µm 

long) ascospores. For differences to T. subadjectum 

see under that species. Thelotrema polythecium is 

distinguished by the absence of the stictic acid chemosydrome, distinctly split ascomata 

margins and a fused proper exciple. Thelotrema subexpallescens from the Andaman Islands 

(and probably Africa, see Frisch, 2006) is similar and might be synonymous with T. 

cyphelloides, type material was unfortunately not available for study. 


Thelotrema defossum (Müll.Arg.) Mangold comb. nov. ined. 

Bas.: Ocellularia defossa Müll. Arg. in Engler, Botan. Jahrbücher 5: 138 (1884). Type: Timor, Mt. 

Taimanani, 1883, Naumann 386 pr. p. (G!-holotype). 



T. cyphelloides. 

Thallus epi- to hypophloedal, predominantly thin, up to c. 200 µm high, pale yellowish 

gray to pale olive. Surface dull to slightly shiny, smooth, continuous to distinctly verrucose, 

distinctly fissured. Thallus covered by an incontinuous protocortex up to 10 µm thick. Algal 

layer well developed, continuous to incontinuous due to crystal inclusions, calcium oxalate 

abundant to sometimes sparse, small to large, clustered or scattered. Vegetative propagules 

not seen. Ascomata inconspicuous, predominantly small, up to c. 300 µm in diam., roundish


Fig. 124. Thelotrema defossum: growth habit (A-C), ascoma section (D) and ascospores (E, F). A., 

B.: A. & K. Kalb 21713; C., E.: G-holotype; D., F.: Lumbsch & Mangold 19161 v. Bar= A: 1 mm; 

B: 0.5 mm; C: 0.75 µm; D: 100 µm; E: 10 µm; F: 8 µm. 

to sometimes slightly elongated, apothecioid, solitary to fused, predominantly immersed, in 

strongly verrucose thalli sometimes becoming somewhat raised. Disc not visible from surface 

to sometimes becoming partly visible, grayish, predominantly epruinose. Pores (moderately) 

small, up to c. 200 µm in diam., roundish to sometimes slightly elongate or somewhat 

irregular, predominantly entire to slightly split, apical proper exciple visible from surface, 

sometimes somewhat shrunken, whitish to off-white, pale brownish towards the base, 

predominantly incurved. Thalline rim margin thin to moderately thick, moderately wide to 

wide, predominantly entire to rarely slightly split, roundish to irregular-roundish to somewhat 

elongate, rarely slightly layered, predominantly incurved to more rarely somewhat erect, 

concolorous with thallus or slightly brighter. Proper exciple becoming partly free in upper 

parts, moderately thin, hyaline internally, pale to dark brown marginally, apically often dark 

brown to slightly carbonized, often amyloid at the base. Hymenium up to c. 80 µm high, noninspersed, 

moderately conglutinated, paraphyses moderately interwoven, unbranched, tips


slightly to moderately thickened, lateral paraphyses present, inconspicuous, up to c. 15 µm 

long, columellar structures absent. Epihymenium indistinct to thin, hyaline, rarely with sparse 

grayish granules. Asci 4-8-spored, tholus moderately thin to moderately thick, thin when 

mature. Ascospores very to moderately small, transversely septate, cell walls predominantly 

(moderately) thin, with distinct, often moderately thick, smooth to irregular halo, hyaline, 

non-amyloid in younger ascospores to weakly amyloid in mature stages, sometimes oblong to 

fusiform to predominantly clavate, ends roundish to subacute, loci roundish to and more often 

±angular, in particular in younger stages, lentiform to flattened cubical, end cells 

hemispherical to conical, septae (moderately) thick with age, regular, 10-30(40) x 5-8 µm 

with 4-11(12) loci. Pycnidia not seen. 


ECOLOGY AND D ISTRIBUTION – Thelotrema 

defossum occurs on tree bark in (sub)tropical to 

warm-temperate rainforests in altitudes ranging 

from sea level to 950 m. This moderately common 

and wide-spread species, occurs in northern 

Queensland, the Queensland/New South Wales 

border region, New South Wales and on Lord 

Howe Island. This is the first report for Australia. 

The species was described from Timor and one 

collection from Florida belongs to this species as 

well (see below). 

NOTES – This species belongs to the T.subtilegroup. 

Within this group of similar species, 

T. defossum is characterized by small, faintly 

amyloid ascospores with thin cell walls, distinct 

halo and flattened-angular loci. The ascospores of 

T. suecicum partly fall within the range of those 

found in T. defossum with 20-40(60) µm in length 

and 6-12(14) loci, but are distinctly broader (8-15 


T. defossum. 

µm), have thick to usually very thick cell walls and different shaped loci (see also under this 

species). Thelotrema parvizebrinum is a similar Australian species, see there for differences. 

One collection from Florida in US determined by Harris as T. lathraeum agrees well with the 

type of T. defossum and the Australian collections. It is therefore possible that T. defossum is 

synonymous to T. lathraeum since it is the older name (Tuckerman, 1882). The type of T. 

lathraeum was unfortunately not available for study. 


19161 s, v, 19162 m, Mangold 31 o (F). Near end of Black Mountain Rd., 33 km WNW of Kuranda, Hale 

832187, 832704 (US). Atherton Tablelands, Lake Euranoo, Lumbsch & Mangold 19127 k (F). Bunya 

Mountains, Mt. Mowbullan, Kalb & Rogers 20283 (hb. Kalb). Wolston nr. Brisbane, Wilson '1464' (as O. cf. 

jugalis) (NSW-539407, US). New South Wales: Nightcap Forest Drive, 1 km W of Minyon Falls, N of Lismore, 

Hale 832087, 832180, 832543 (US). Iluka Nature Reserve, 50 km NE of Grafton, Mangold 23 i (F). Blue Mnts., 

Below Bridal Veil Falls, near Blackheath, Hale 58543 (US). Royal NP., S of Sydney, Bola Creek, E of 

Waterfall, K. & A. Kalb 21711, 21713 (hb. Kalb). Lord Howe Island, near Golf Club, Elix 42072 (CANB). 

U.S.A., Florida, Hale 53103 (US).


Thelotrema diplotrema Nyl. 

Ann. Sci. Nat., Bot., sér. 4, 11: 258 (1859). Ocellularia diplotrema (Nyl.) Zahlbr., Cat. Lich. Univ. II: 588 

(1923). Type: Réunion ['Borbonia'], s.c. (H-Nyl. 22737!-lectotype, selected by Hale [1972 in herb.]). 

Ocellularia turgidula Müll. Arg., Journ. de Botan. 7: 94 (1893). Thelotrema turgidulum (Müll. Arg.) Hale, 

Mycotaxon 11: 132 (1980), nom. illeg. [non Thelotrema turgidulum Vain.]. Type: New Caledonia, Mt. Mou, 

1870, Balansa s.n. (PC-lectotype, selected by Hale [1978: 33]; G!-isolectotype). 


Fig. 126. Thelotrema diplotrema: growth habit (A), ascomata (B, C), ascoma section (D), ascospore 

(E), ascospores showing amyloid reaction (F) and ascospore detail (G). A., E.-G.: Lumbsch & 

Mangold 19127 v; B.: G-isolectotype of O. turgidula; C.: H-lectotype; D.: Lumbsch & Mangold 

19139 r. Bar= A: 1 mm; B: 0.35 mm; C: 0.75 mm; D: 100 µm; E: 11 µm; F: 9 µm, G: 7 µm.


Thallus epi- to hypophloedal, moderately thin, up to c. 200 µm high, pale greenish gray to 

pale olive. Surface predominantly dull to rarely slightly shiny, smooth to more often 

roughened, sometimes porous, continuous to ±distinctly verrucose to verruculose, slightly to 

distinctly fissured. Cortex structures absent or covered by a usually thin, incontinuous 

protocortex up to c. 20 µm thick. Algal layer well developed, continuous to incontinuous, 

often due to calcium oxalate crystal inclusions, calcium oxalate crystals abundant, 

predominantly small, often clustered. Vegetative propagules not seen. Ascomata usually 

inconspicuous, predominantly small to rarely moderately large, up to c. 400(700) µm in 

diam., roundish to slightly irregular, apothecioid, solitary to marginally or entirely fused, 

immersed to ±distinctly emergent, then flattened-hemispherical to flattened-urceolate, with 

same surface as thallus. Disc often becoming partly visible from surface, (pale) grayish to 

pale flesh-colored, epruinose to slightly pruinose. Pores (moderately) small, up to c. 150 µm 

in diam., roundish to slightly irregular, entire to split, apical to rarely entire proper exciple 

visible from surface, often only slightly separated from thalline rim margin, rarely somewhat 

shrunken, whitish to off-white, pale brownish towards the base, incurved to somewhat erect. 

Thalline rim margin moderately thin to more often moderately thick, predominantly entire, 

roundish to irregular-roundish, becoming moderately wide, incurved to more rarely somewhat 

erect, concolorous with thallus to sometimes brownish. Proper exciple becoming apically to 

partly, rarely entirely free, moderately thin to moderately thick, hyaline to pale yellowish 

internally, brownish or yellowish-brown marginally, apically often dark-brown, often amyloid 

at the base. Hymenium up to c. 180 µm high, non-inspersed, moderately conglutinated, 

paraphyses ±interwoven, unbranched, tips moderately to distinctly thickened, lateral 

paraphyses present, predominantly inconspicuous, up to c. 30 µm long, columellar structures 

absent. Epihymenium variable, indistinct to moderately thick, hyaline, with grayish to 

sometimes brownish granules and small crystals. Asci 4-8-spored, tholus (moderately) thick, 

thin when mature. Ascospores moderately large, transversely septate, very rarely with a single 

longitudinal septum, cell walls thick, often slightly crenate, thinly halonate, hyaline, distinctly 

to strongly amyloid, sometimes clavate to more often ±fusiform, ends roundish to narrowedroundish 

to more rarely subacute, loci ±roundish, subglobose to lentiform or somewhat 

irregular, end cells hemispherical to conical, septae (moderately) thick, regular to slightly 

irregular, 50-90(110) x 8-12 µm with 14-20(22) (x 2) loci. Pycnidia not seen. 




diplotrema was collected in Australia on tree bark 

predominantly in (sub)tropical to warm-temperate 

rainforests, more rarely in wet sclerophyll forests in 

altitudes ranging from sea level to 1100 m. It is a 

common and widespread species occurring in 

Queensland and northern New South Wales. It is 

also known from Central America (Hale, 1978), 

Africa (Frisch, 2006) Reunion, Andaman Islands 

(Sethy & al., 1987) and Japan (Hale, 1978), 

indicating a pan(sub)tropical distribution. 


ecorticate, often roughened thallus, immersed to 

emergent, small apothecia with often only 

indistinctly free proper exciple, moderately large, 


T. diplotrema.


transversely septate, thick-walled, distinctly amyloid ascospores, and the absence of 

secondary compounds. It is similar to T. pseudosubtile, especially when younger or poorly 

developed ascomata are observed that lack fully developed ascospores; see under this species 

for differences. Thelotrema diplotrema is another member of the T. subtile-group. Within this 

group T. nureliyum is also similar but readily distinguished by larger ascomata with a more 

distinctly free proper exciple and larger ascospores (up to 220 µm long with up to 35 loci). As 

mentioned above, T. bicavatum was included as a (tentative) synonym to T. diplotrema 

(Frisch, 2006). This species, however, differs in smaller ascospores (up to 45 µm long with up 

to 16 loci) and contains norstictic acid and is therefore here regarded as a distinct taxon. 


19161 r (F). Mt. Windsor, 5 km W of new Forestry Camp, NW of Mossman, Hale 830868 (US). Near Lake 

Placid, S of Kuranda, K. & A. Kalb 21301 (hb. Kalb). End of Clohesy River Rd., 16 km SE Kennedy Hwy., W 

of Cairns, Hale 832241 (US). Atherton Tablelands: Davies Creek Rd. 17 km S of Kennedy Hwy., S of Davies 

Creek Falls NP., E of Mareeba, Hale 831247, 832691 (US); Danbulla Forest Drive: 4 km E of Tinaroo, Hale 

831210 (US), Lake Euramoo, Hale 831521 (US), Lumbsch & Mangold 19127 v (F); Area below crater, Mt. 

Hypipamee NP., S of Atherton, Hale 832550 (US); Millaa Millaa falls, Lumbsch & Mangold 19139 j, r (F); 

Souita Falls, Lumbsch & Mangold 19155 y (F); Along road going S from Ravenshoe to Tully Falls, Hale 832566 

(US); 13 km S of Ravenshoe on Tully Falls Rd., Hale 831286, 831696 (US); Culpa logging area, 13 km from 

Koombooloomba rd. turnoff, SE of Tully Falls, Hale 832119 (US). Murray Falls, W of Kennedy, Hale 831612, 

832465 (US). Mt. Spec NP., Ridge on the Loop, on the Paluma Rd., WNW of Townsville, Hale 832321, 832760 

(US). Conway Range NP., near Shute Harbour-Airlie Beach, Hale 830882 (US). Eungella NP.: 'Credition 

Track', K. & A. Kalb 26678 (hb. Kalb); Rosser Rd. entry point off Darymple rd. (near Pease's Lookout), Hale 

831126, 831351, 381354, 831623, 831406, 831745 (US). Mt.Mee SF., 6 km NW of Forestry Office, NW of 

Mt.Mee, Hale 832553, 832782 (US). D'Anguilar Range NW of Brisbane, about 2 km N of Mt.Glorious W of the 

rd. to Mt.Tenison Woods, Hafellner 16988 (GZU). Brisbane, [as T. rimulosum], Bailey s.n. (NSW-539340). 

Cunninghams Gap NP., 50 km NE of Warwick, Hale 831733 (US). Carabeen Nature Refuge, 45 km E of 

Warwick, Lumbsch & Mangold 19174 m, 19175 b (F). Best of All Lookout, 4 km S of Springbrook, Wunburra 

Range, Elix 2517 (CANB). Lamington NP.: Python Rock Track, Hale 832736, 832755 (US); Main Border Track 

out of O'Reillys, Hale 830962, 831930 (US). New South Wales: Lions Tourist Rd. near Queensland border, N of 

Waingaree, Hale 830942 (US). Tweed Range, Mebbin NP., 25 km SW of Murwillumbah, Mangold 21 c, d, h, j, 

k (F). Gibbergunyah Roadside Reserve, Nightcap Forest Drive, Whian Whian SF., W of Mullumbimby, Hale 

832538 (US). Nightcap NP., Mnt. Nardi/Mnt. Matheson Track, Mangold 22 h, i (F). Waingaree Forest, N of 

Kyogle: Black Butt Picnic Area, Loop Drive, Hale 830285 (US); NW part of Wiangaree Forest Drive, Hale 

831192, 831390, 831391, 831524, 831616, 832471, 832502 (US). Mt.Warning NP., track from summit to 

parking lot, Mangold 19 d, l, p, q, r, y, za, zf, zi (F). Cambridge Plateau Forest Drive, 3 km N of picnic area, 

Richmond Range SF., 30 km W of Casino, Hale 832486 (US). Richmond River, Jun. 1884, Wilson s.n. (NSW- 

539420). Dorrigo NP.: Never Never Picnic Area and Rosewood Creek Track, Mangold 24 a, g, h (F); Sassafras 

Creek Track, Mangold 25 e, t (F). Doyles River SF. on Oxley Hwy., 95 km SE Walcha, Hale 58590 (US). 

Barrington Tops NP., NE of Scone, K. & A. Kalb 21787 (hb. Kalb). Bulahdelah District, Myall River SF., E of 

Stroud, Jarrah Rd., Kalb & Filson 17888, 17940, 17949, 17955, 17960 (hb. Kalb). Royal NP., S of Sydney, Bola 

Creek, E of Waterfall, K. & A. Kalb 21712, 21722 (hb. Kalb). Monga NP., 27 km SE of Braidwood, Mangold 11 

w (F). New Caledonia: Riviere Blanche, Hill 11716 (US); Yate, Hill 11768 (US). 

Thelotrema eungellaense Mangold, Elix & Lumbsch 

Bibl. Lichenol. 95: 463 (2007). Type: Australia, Queensland, Eungella NP., trail from Broken River Station 

to rainforest, Lumbsch & Mangold 19108 k (CANB!-holotype, BRI!-isotype). 


Thallus hypo- to epiphloedal, thin to moderately thick, up to c. 400 µm high, pale grayishor 

yellowish-green to olive. Surface dull, smooth, ±verrucose, fissured. True cortex present, 

continuous to sometimes incontinuous, up to c. 50 µm thick, consisting of irregular to more 

often periclinal hyphae. Algal layer variable, continuous to incontinuous, moderately well to 

poorly developed, calcium oxalate crystals abundant, small to large, scattered to clustered. 

Vegetative propagules not seen. Ascomata usually conspicuous, moderately large, up to c.


800 µm in diam., roundish, perithecioid, solitary, immersed to emergent, then predominantly 

(verrucose-)hemispherical to more rarely (verrucose-)subglobose. Discs not visible from 

surface. Pores small, up to c. 150 µm in diam., roundish, entire, apical proper exciple usually 

becoming visible from surface, forming a predominantly entirely free inner pore margin, 

often slightly sunken, incurved, whitish to off-white. Thalline rim margin moderately thick to 

more often conspicuously thick, roundish, entire to more rarely slightly split, usually whitish 

to off-white or brighter then thallus, sometimes concolorous with thallus or more rarely 

brownish, level with thalline rim to ±distinctly annulate or funnel-shaped to more rarely 

sunken, thalline rim not developed or not distinguishable from thallus in immersed ascomata, 

in emergent ascomata incurved, with same surface as thallus. Proper exciple partly fused to 

entirely free, moderately thick, hyaline to pale yellowish internally, yellowish-gray 

marginally, apically sometimes brownish, non-amyloid to more rarely faintly amyloid at the 


thin, distinctly bent to curly towards tips, parallel to slightly interwoven, unbranched, tips 

slightly thickened, lateral paraphyses present, mostly inconspicuous, not conglutinated, up to 

c. 40 µm long, columellar structures absent. Epihymenium indistinct, hyaline, without 

granules or crystals. Asci 2-4-spored, tholus thick, thin when mature. Ascospores large, 

densely eumuriform, cell walls and endospore thin, non-halonate, hyaline, rarely pale 

yellowish at late maturity, non-amyloid to faintly amyloid in mature spores, predominantly 

ellipsoid to fusiform, with narrowed-roundish to rarely slightly subacute ends, rarely slightly 

bent, loci small, roundish to slightly angular, subglobose to roundish-cuboid or irregular, 

transverse septae thin, distinct throughout development, regular to slightly irregular, 80-180 x 

20-40 µm, with multiple loci. Pycnidia not seen. 

Fig. 128. Thelotrema eungellaense: growth habit (A), ascomata (B) and asccospores (C, D). A.: Hale 

832175; B., D.: CANB-holotype; C.: Hale 831311. Bar= A: 1 mm; B: 0.65 mm; C: 25 µm; D: 25 

µm.





eungellaense grows on tree bark in tropical 

highland rainforests in altitudes ranging from 600- 

1500 m. It is moderately common in northern 

Queensland and is currently only known from 

there. 


verrucose, corticate thallus, perithecioid ascomata 

with a ±free proper exciple, large, densely 

eumuriform, hyaline, thin-walled, non-amyloid or 

indistinctly amyloid ascospores, and the norstictic 

acid chemosyndrome. It is similar to T. porinaceum, 

which differs by larger ascospores (up to 230 

µm long). Another similar species is the 

paleotropical T. weberi, this taxon differs by a thin, 

evanescent thallus and large, smooth, urceolate, 

wide-pored ascomata and distinctly amyloid, thickwalled 

ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Mt. Windsor Rd., NW of Mossman, Hale 831310, 831311 

(US). Mt. Windsor logging area, NW of Mossman, Hale 830842, 832175 (US). Mt. Lewis Rd., W of Mossman, 

Hale 830625, 830643, 832172, 832399, 832620, 832659, 832661 (US). Davies Creek Rd., S of Davies Creek 

Falls NP., Hale 830662, 830631, 830864 (US). Atherton Tablelands: Plath Rd. logging head, S of Atherton, 

Hale 830649, 831845, 831944, 832641 (US); Mt. Hypipamee NP., Hale 830646, 832814 (US); Tumoulin Rd., 5 

km from turnoff to Ravenshoe, Mangold 30 y (F); 10 km S of Ravenshoe on Tully Falls Rd., Hale 831016 (US). 

Bellenden Ker NP., centre peak, A. & M. Aptroot 22484, 22512 (ABL). Dawson logging area, WSW of Tully, 

Hale 830698, 830905, 830906, 832136, 832520, 832660, 832663, 832693 (US). Culpa Logging area, SE of 

Tully Falls, Hale 830628, 830703, 832190, 832137, 832584, 832652 (US). Mt. Spec NP., WNW of Townsville, 

Hale 830632, 830647, 830648, 830659, 830722, 80915, 831608, 832134, 832545, 832774 (US). Eungella NP.: 

Rd. to rainforest at NP. margin, Lumbsch & Mangold 19110 d (F); Rosser Rd. entry point off Darymple rd., Hale 

831525, 831740 (US). 

Thelotrema foveolore Müll. Arg. 

Nuov. Giorn. Bot. Ital. 23: 129 (1891). Type: Japan, Tosa, (com. Miyoshi 110), Yoshinaza 26 (G!-holotype). 



T. eungellaense. 

Thallus epi- to hypophloedal, (moderately) thin, up to c. 150 µm high, grayish to 

yellowish- or greenish-gray. Surface dull, smooth, continuous to verrucose, fissured to 

sometimes appearing areolate due to substrate structure. Cortex structures absent or with a 

usually thin, incontinuous protocortex up to c. 20 µm thick. Algal layer poorly developed, 

incontinuous, calcium oxalate usually abundant in ascomata area, otherwise sparse, small to 

large, clustered. Vegetative propagules not seen. Ascomata conspicuous, large, up to 1.2 mm 

in diam., roundish to slightly irregular, perithecioid when young, becoming apothecioid at 

maturity, solitary to marginally fused, ±distinctly emergent, predominantly (irregular- 

)subglobose to (irregular-)urceolate, sometimes flattened-cylindrical. Disc not visible from 

surface. Pores opening in late stages, relatively small, up to c. 400 µm, roundish to more often 

roundish-irregular to irregular, apical proper exciple visible from surface, becoming entirely


Fig. 130. Thelotrema foveolare: growth habit (A, B), ascoma section (C), asci (D, E) and ascospore 

detail (F). A., C., D.: Hale 831230; B.: G-holotype; E., F.: Hale 83146. Bar= A: 1.5 mm; B: 2 mm; 

C: 200 µm; D: 10 µm; E: 25 µm; F: 10 µm. 

free, off-white, often shrunken, incurved to somewhat erect. Thalline rim margin roundish to 

irregular, (moderately) thick, wide to gaping, predominantly entire, incurved, concolorous 

with thallus to brownish due to protuberant substrate. Proper exciple becoming entirely free, 

moderately thick, hyaline internally, (pale) brownish to brownish-gray or grayish marginally, 

apically often covered by grayish granules, non-amyloid. Hymenium up to c. 300 µm high, 

non-inspersed, moderately conglutinated, paraphyses parallel or slightly interwoven, 

unbranched, tips moderately to distinctly thickened, lateral paraphyses present, 

inconspicuous, up to c. 20 µm long, columellar structures absent. Epihymenium variably 

thick, indistinct in younger stages to moderately thick in older ascomata, hyaline, with fine to 

coarse grayish or grayish-brown granules. Asci 1-spored, tholus absent, lateral ascus walls 

distinctly thickened, becoming thin when mature in lower parts. Ascospores (very) large, 

densely eumuriform, in early stages cell walls and endospore distinctly thickened, becoming 

moderately thin to moderately thick at maturity, non-halonate, hyaline to rarely somewhat 

yellowish in depauperate ascospores, loci strongly to opaque amyloid, cell walls and 

endospore non-amyloid, predominantly fusiform, with narrowed-roundish to subacute ends, 

loci large, roundish to slightly angular, subglobular to roundish-cuboid, transverse septae


(moderately) thin, distinct, regular, 110-230(250) x 20-40(50) µm with multiple loci. Pycnidia 

not seen. 

CHEMISTRY – Thallus K+ yellowish, C-, PD+ yellow; containing psoromic and 2'-0demethylpsoromic 

acids (majors). 


foveolare was collected in Australia on tree bark in 

tropical highland rainforests in altitudes ranging 

from 530 to 1200 m. It is a moderately common 

species occurring in northern and north-central 

Queensland. This is the first report for Australia, it 

is otherwise only known from Japan. 

NOTES – Thelotrema foveolare is a uniform 

species, which is characterized by large, emergent 

ascomata with relatively small pores and free 

proper exciple, monosporic asci that lack a tholus 

but have distinctly thickened lateral walls, large, 

densely eumuriform, hyaline, ±thick-walled, 

amyloid ascospores, and the presence of the 

psoromic acid chemosyndrome. Ascospores with a 

similar reaction in iodine - the inner parts turning 

distinctly purple, cell walls and endospore remaining 

hyaline - can be found in Topeliopsis 

azorica and Thelotrema laceratulum, two taxa that 


T. foveolare. 

otherwise differ in several characters (ascomata with ±lacerate thalline rim margins, 

ascospores not exceeding 160 µm in length, containing either stictic acid or lacking secondary 

compounds). Similar is T. saxicola, which can be readily distinguished by fused proper 

exciple and an inspersed hymenium. Similar species in Australia with different secondary 

chemistry are T. porinaceum and T. rugatulum. Thelotrema porinaceum shares the same 

ascus-type with T. foveolare but contains norstictic acid and is further distinguished by 

perithecioid ascomata at maturity. Thelotrema rugatulum lacks lichen substances, has asci 

with distinct tholus, at least in younger stages, and further differs in a slightly lower 

hymenium (up to 250 µm high) and ascospores without distinctly thickened walls. The type 

collection differs from Australian material slighly in having indistinctly emergent ascomata 

with tiny ostioles and resembles an immature sample. One Hale collection from Townsville 

area (Hale 831887) differs in less distinctly emergent ascomata, 8-spored asci with non- or 

faintly amyloid ascospores. This specimen is tentatively included here. 

SPECIMENS EXAMINED – Australia, Queensland: Mt.Windsor logging area, near jct. rd. to old Forestry Camp 

and main rd., NW of Mossman, Hale 831902, 832621, 832696 (US). Mt.Windsor, 5 km W of new Forestry 

Camp, NW of Mossman, Hale 832688 (US). End of Clohesy River Rd. 16 km SE Kennedy Hwy., W of Cairns, 

Hale 830835, 830879, 831900 (US). Atherton Tablelands: Lamb Range, near Mt. Haig, 23 km SE of Mareeba, 

Streimann 57728 c (B); Area below crater, Mt. Hypipamee NP., Dinner Creek Falls, S of Atherton, Hale 

831392, 831469, 832339 (US); SW of K-1 tree rd. off Palmerston Hwy., 11 km from main hwy. and 2 km N of 

S. Johnstone Forestry Camp, SE of Millaa Millaa, Hale 830859 (US); Just S of hwy., 13-23 km E of jct. 

Kennedy Hwy. and Palmerston Hwy., E of Ravenshoe, Hale 831277, 831284, 831908, 832176, 832511 (US). 

About 7.5 km E of Wallaman Falls, W of Ingham, Hale 832245 (US). Mt.Spec NP., Ridge on the Loop, on the 

Paluma Rd., WNW of Townsville, Hale 831887, 831921 (US). Cedar Creek Falls, SE of Mt.Julian, 33 km S of 

Airlie Beach, Hale 831124 (US). Eungella NP., NP. side rd. nr. Peases Lookout, off Darymple rd., Hale 830749, 

831230, 831233, 831551, 831647, 831711 (US).


Thelotrema gallowayanum Mangold, Elix & Lumbsch 

Bibl. Lichenol. 95: 465 (2007). Type: Australia, Queensland, Atherton Tablelands, Tumoulin Rd., 5 km from 

turnoff to Ravenshoe, Lumbsch & Mangold 19151 t (CANB!-holotype; BRI!-isotype). 


Fig. 132. Thelotrema gallowayanum: growth habit (A), ascoma with emerging ascospore (B), ascoma 

section (C), ascospore (D) and details of ascospores (E, F). A.-F.: CANB-holotype. Bar= A: 1.5 

mm; B: 0.2 mm; C: 0.25 mm; D: 50 µm; E: 25 µm; F: 15 µm. 

Thallus epi- to hypophloedal, thin to moderately thick, up to c. 400 µm high, pale grayishgreen 

to pale yellowish-gray. Surface dull, smooth to roughened, continuous to more often 

verruculose to verrucose, fissured. Thallus covered by an incontinuous protocortex, up to c. 

20 µm thick. Algal layer variable, continuous to incontinuous, moderately well to poorly 

developed, calcium oxalate crystals abundant, small to large, scattered to clustered. 

Vegetative propagules not seen. Ascomata inconspicuous, moderately large, up to c. 800 µm 

in diam., roundish, perithecioid, solitary, immersed to emergent, then predominantly 

hemispherical. Disc not visible from surface. Pores tiny to small, up to c. 120 µm in diam., 

roundish, entire, apical proper exciple sometimes becoming visible from surface, then 

forming a fused to more often entirely free inner pore margin, often ±sunken, incurved, 

predominantly off-white, otherwise pore margin formed by thalline rim. Thalline rim margin 

moderately thick, roundish, entire, concolorous with thallus to whitish or brownish, level with


thalline rim, thalline rim not distinguishable from thallus in immersed ascomata, in emergent 

ascomata incurved, with same surface as thallus. Proper exciple partly fused to entirely free, 

thin, hyaline to pale yellowish internally, yellowish-gray marginally, non-amyloid. 

Hymenium up to c. 500 µm high, non-inspersed, moderately conglutinated, paraphyses thin, 

distinctly bent, parallel to slightly interwoven, unbranched, tips slightly thickened, lateral 

paraphyses present, inconspicuous, not conglutinated, up to c. 30 µm long, columellar 

structures absent. Epihymenium indistinct, hyaline, without granules or crystals. Asci 1- to 

very rarely 2-spored, tholus thick, moderately thin when mature. Ascospores typical, very 

large, densely eumuriform, cell walls and endospore (moderately) thin, non-halonate, brown 

to yellowish-brown at late stage of maturity, non-amyloid to faintly amyloid, predominantly 

oblong-fusiform, sometimes slightly irregular in outline, straight to slightly bent, ends typical, 

strongly tapered, forming distinct short conical appendixes with 3 - 6 transversely septate loci, 

loci roundish to angular, particularly in later stages, subglobose to oblong or irregular, 

transverse septae thin, remaining distinct and regular throughout development, 180-350(400) 

x 20-45(50) µm, with multiple loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ orange-red, C-, PD-; containing norstictic (major) and connorstictic 

(minor to trace) acids. 


gallowayanum was grows on tree bark in 


sea level to 1100 m. It is moderately common and 

wide-spread in Queensland and the Queensland/New 

South Wales border region. So far it is 

only known from there. 

NOTES – This taxon can be easily identified by 

the inconspicuous, perithecioid ascomata, large, 

densely eumuriform, thin-walled, brown ascospores 

with tapered ends and the presence of 

norstictic acid. Thelotrema eungellaense and 

T. porinoides, both also with norstictic acid, are 

similar species in Australia, both readily 

distinguished by smaller, hyaline ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Daintree 

NP., Mossman sct., lower Mossman River, Mangold 36 m (F). 

Mt.Windsor, 5 km W of new Forestry Camp, Hale 830618 


T. gallowayanum. 

(US). Mt.Lewis Rd., 13 km N from Kennedy Hwy., Hale 831237 (US). Davies Creek Rd., S of Davies Creek 

Falls NP., Hale 830856 (US). Atherton Tablelands: SW of K-1 tree rd. off Palmerston Hwy., Hale 832331 (US); 

Lake Barrine NP., Hale 830756 (US); Mt.Hypipamee NP., Hale 830627 (US); Tumoulin Rd., 5 km from turnoff 

to Ravenshoe, Lumbsch & Mangold 19151 b, f, g, r, s, zb, zc (F); Mangold 30 r, y (F). Kalpowar Forest Drive, 

40 km NE of Monto, Hale 831712 (US). Bunya Mnts. NP., Hale 831486 (US). Wooroi SF., W of Tewantin, 

Hale 830655 (US). Mt.Spec NP., Paluma Rd., Hale 830907 (US). Mt.Mee SF., 6 km NW of Forestry Office, 

Hale 830844, 832727 (US). Lamington NP.: A. & M. Aptroot 21834 (ABL); Tibell 12713 (UPS). New South 

Wales: Iluka Nature Preserve: Mangold 23 g (F); Hale 58718, 58734 (US).


Thelotrema lacteum Kremp. 

in Nyl., Flora 47: 269 (1864). Phaeotrema lacteum (Kremp.) Müll.Arg., Flora 70: 398 (1887). Type: 

Australia, com. Hochstetter (M!-lectotype, selected by Hale [1981: 256]; H-Nyl. 22814!- isolectotype). 

Phaeotrema consimile Müll.Arg., Flora 70: 398 (1887). Type: Australia, Queensland, Russell River, Sayer 

s.n. (G!-lectotype, here selected). 

Thelotrema sitianum Vain., Étud. Lich. Brésil 2: 81 (1890). Phaeotrema sitianum (Vain.) Zahlbr., in Engler- 

Prantl, Nat. Pflanzenfam. 1: 119 (1905). Thelotrema lepadodes Tuck. ssp. sitianum (Vain.) Salisb., 

Lichenologist 5: 270 (1972). Type: Brazil, Minas Gerais, Vainio - Lich. bras. exs. 565 (TUR-Vain. 26808!lectotype, 

selected by Salisbury [1972: 270]). 

Ocellularia cricota F.Wilson in Bailey, Bot. Bull. Dept. Agric., Queensland 7: 32 (1891). Phaeotrema 

cricotum (F.Wilson) Müll.Arg., Hedwigia 32: 130 (1893). Type: Australia, Queensland, Southport, Wilson s.n. 

(G-lectotype, selected by Hale 1981: 256] not seen; BRI-'Shirley Book', p. 22, n. 11 [BRI-AQ721231]!isolectotype]). 

Ocellularia zeorina Müll.Arg., Nuovo Giorn. Bot. Ital. 23: 394 (1891). Type: Australia, Brisbane, Bailey 460 

pr. p. [with O. profunda] (G!-holotype). [The material in BRI-'Shirley Book', p. 22 notated as O. zeorina is the in 

Müller's description mentioned intermixed specimen determined as Campylothelium nitidum Müll.Arg.]. 

Ocellularia annulosa Müll.Arg., Bull. Herb. Boissier 3: 314 (1895). Type: Australia, Queensland, [app. Mt. 

Gravatt (Brisbane)], 1894, Shirley s.n. (G!-holotype; BRI-'Shirley Book', p. 21 [BRI-AQ721217]!-isotype). 


Fig. 134. Thelotrema lacteum: growth habit (A, B), ascomata (C, D), ascoma section (C), younger 

ascospores (F) and mature ascospores (G). A.: M-lectotype; B., C.: Mangold 27 p; D.: Hale 

34276; E.: H-isolectotype; F.: Hafellner 19629; G.: TUR-lectotype of T. sitianum. Bar= A: 1 mm; 

B: 1.25 mm; C: 1.25 mm; D: 0.75 mm; E: 200 µm; F; 20 µm; G: 25 µm.


Thallus variable, thin to moderately thick, epi- to hypophloedal, up to c. 300 µm thick, pale 

gray to pale yellowish-gray or pale grayish-green. Surface dull to sometimes slightly shiny, 

smooth to rough, continuous to verrucose or verruculose, often ± fissured. Thallus 

predominantly covered by an ±continuous protocortex up to c. 30 µm thick, rarely becoming 

distinctly conglutinated forming a true cortex of periclinal hyphae. Algal layer variable, 

continuous to more often discontinuous, well to poorly developed, calcium oxalate crystals 

usually abundant, small to large, scattered or in clusters, sometimes sparse. Vegetative 

propagules not seen. Ascomata usually conspicuous, (moderately) large, up to c. 1 mm in 

diam., roundish to slightly irregular, apothecioid to more rarely somewhat chroodiscoid, 

somewhat erumpent to sessile, solitary to fused, ±distinctly emergent, predominantly 

(depressed-)urceolate to depressed cylindrical. Disc usually partly to entirely visible from 

surface, pale grayish to usually whitish, distinctly to strongly pruinose. Pores moderately wide 

to gaping, up to c. 800 µm in diam., roundish to (slightly) irregular, apical to upper proper 

exciple visible from surface, free, predominantly entire and ±jagged, sometimes eroded, 

whitish to off-white, often pruinose, incurved to more often erect or recurved, sometimes 

slightly shrunken. Thalline rim margin predominantly thick, roundish to slightly irregular, 

entire to more often distinctly split to lacerate, often eroded, concolorous with thallus, thalline 

rim concolorous and with same surface as thallus, predominantly erect to slightly recurved. 

Proper exciple free, (moderately) thick, pale yellowish internally, yellowish-brown to grayishbrown 

marginally, apically sometimes dark brownish and usually covered by grayish 

granules, ±distinctly amyloid at the base. Hymenium up to c. 200 µm high, non-inspersed, 

moderately conglutinated, paraphyses straight to slightly bent, parallel to slightly interwoven, 

unbranched, tips slightly thickened, somewhat irregular, lateral paraphyses present, 

sometimes inconspicuous, up to c. 30 µm long, columellar structures absent. Epihymenium 

predominantly thick, hyaline, with dark gray granules. Asci 4-8-spored, tholus thick, 

moderately thin when mature. Ascospores (moderately) large, transversely septate, cell walls 

moderately thick to very thick, becoming distinctly crenate in older stages, in younger stages 

with thick, irregular halo, hyaline, becoming distinctly brown usually in later stages of 

maturity or when decayed, moderately to strongly amyloid, oblong to oblong-fusi- to 

claviform with rounded to subacute ends, often ±bent, loci roundish to slightly angular, 

subglobose to lentiform, end cells hemispherical to conical, septae moderately thin to 

moderately thick, regular, (50)70-110(130) x 8-12 µm with (12)16-24(26) loci. Pycnidia not 

seen. 




lacteum grows on tree bark in mangroves, beech 

forests, wet sclerophyll forest and rain forests in 

(sub)tropical to warm-temperate climates in 

altitudes ranging from sea level to 920 m. It is 

moderately common and wide-spread, occurring in 

northern and southern Queensland and northern 

and south-central New South Wales. It is also 

known from Brazil and Japan (Hale, 1981). 

NOTES – The ascomata of this taxon are almost 

Chapsa-like with exposed and conspicuously 

bright discs and a erect to recurved proper exciple. 

It is further characterized by a pale, ecorticate to 


T. lacteum.


indistinctly corticate thallus, moderately large, transversely septate, narrow, thick-walled, 

indistinctly brown, amyloid ascospores and the absence of secondary com-pounds. A similar 

species is T. pachysporum, which has smaller (up to 75 µm long), broader ascospores. 

Specimens with young, unpigmented ascospores may be confused with T. diplotrema, which 

differs in having constantly hyaline ascospores and ascomata with darker, less distinctly 

pruinose discs and incurved to slightly erect, but never recurved proper exciple. The type 

specimen of Phaeotrema consimile is in poor condition and has ascomata with small, entire, 

almost round and slightly brown rimmed margins with hardly visible proper exciples. Since 

anatomically identical to the other examined collections, it is considered conspecific. 

Salisbury (1972) mistakenly identified T. lepadodes (= T. monosporum) with transverse 

septate ascospores (see also under remarks for T. pachysporum) and combined T. sitianum as 

a subspecies of T. lepadodes, differing in larger ascospores. The examination of the type 

revealed that it is conspecific with T. lacteum, slightly differing in somewhat larger 

ascospores. One collection from southern Queensland (Rogers 2604) has small-pored 

ascomata with an incurved exciple and ascospores that are consistently slightly above average 

(up to 130 µm) although the number of loci is in accordance with those usually found in T. 

lacteum (up to 26). 

SPECIMENS EXAMINED – Australia, Queensland: Atherton Tablelands, Dunbulla Forest Drive, near Lake 

Euranoo parking lot, Mangold 37 g (F). Noosa River NE of Tewantin, about 70 km SE of Gympie, Hafellner & 

Stevens 19629 (GZU). E of the Tingarana Lookout SE of Noosa Heads, Hafellner & Stevens 16832 (GZU). 

Bunya Mountains, rd. from the ridge to Maidenwell, Hafellner & Rogers 19688 (GZU). Mt. Tennison Woods, 

near Mt. Glorious, Rogers 2604 (BRI). Moggill SF., Mt. Crosby, Brisbane, Stevens 694628 (BRI). New South 

Wales: Watercatchment of the Karuah River, W of Stroud, Filson 15317 b (MEL). Myall Lakes NP., Mungo 

Brush Camping Area, Mangold 27 a, h, n, p, y, z, ze (F). Trail on eastern bank of Berowra Creek, 25 km NNW 

of Sydney, K. & A. Kalb 34274, 34276 (hb. Kalb). 

Thelotrema lepadinum (Ach.) Ach. 

Meth. Lich.: 132 (1803). Bas.: Lichen lepadinus Ach., Lich. Suec. Prodr.: 30 (1798). Type: Sweden, 

Ostrogathia, Acharius s.n. (UPS-Ach.-lectotype, selected by Purvis & al. [1995: 344]). 

Thelotrema aemulans Kremp., Verhandl. zool.-bot. Gesellsch. Wien 26: 453 (1876). Leptotrema aemulans 

(Kremp.) Müll. Arg., Bull. Herb. Boissier 2(1): 75 (1894). Type: New Zealand, hb. Krempelhuber n. 12, Knight 

s.n. (M!-holotype). 

Thelotrema flavescens Darb. in Nordenskjöld, Wiss. Ergebnisse Schwed. Südpolarexpedit. 1901-1903, 4(4): 

6 (1912). Type: Chile, Tierra del Fuego, Navarin Island, 04. Mar. 1902, Skottsberg s.n. (S!-holotype). 

Thelotrema obconicum Raes., Arch. Soc. zool.-bot. fenn. Vanamo 3: 184 (1949). Type: Australia, New South 

Wales, Katoomba, Sept. 1889, Wilson s.n. (H!-holotype). 

[For additional synonyms see Frisch 2006, Salisbury 1972 and Zahlbruckner 1923.] 



brown to pale grayish-green or pale olive. Surface dull to slightly shiny, smooth, continuous 

to distinctly verrucose, slightly to distinctly fissured. Cortex structures variable, 

predominantly an incontinuous protocortex present, up to c. 30 µm thick, in some parts, 

usually at thalline rim, becoming distinctly conglutinated forming a true cortex of irregular to 

periclinal hyphae. Algal layer usually ±well developed and continuous, often becoming 

somewhat incontinuous due to calcium oxalate crystal inclusions, calcium oxalate crystals 

±abundant, small to large, scattered to clustered. Vegetative propagules not seen. Ascomata 

usually conspicuous, moderately large to very large, up to c. 2 mm in diam., roundish to 

sometimes slightly irregular, apothecioid, sessile, solitary to sometimes marginally fused, 

distinctly emergent, hemispherical to more often urceolate or subglobose, with same surface 

as thallus to often more distinctly shiny. Disc sometimes becoming partly visible from


Fig. 136. Thelotrema lepadinum: growth habit (A), ascomata (B), ascoma and thallus section (C), 

pycnidia (D), condidia (E), ascospores (F) and same ascospores shwowing amyloid reaction (G). 

A.: Verdon 3882; B.-G.: CANB-holotype. Bar= A: 1.5 mm; B: 1 mm; C: 150 µm; D: 0.1 mm; E: 5 

µm; F, G: 4 µm. 

surface, grayish to brownish, indistinctly pruinose. Pores small to moderately wide, up to c. 

500 µm in diam., roundish to more often irregular, entire to split, apical to more often entire 

proper exciple visible from surface, free, usually shrunken, apically bright, bright yellowish 

towards the base, incurved. Thalline rim margin thin to thick, entire to somewhat eroded, 

±roundish to more rarely slightly elongate, usually becoming wide to gaping, incurved to 

somewhat erect, predominantly concolorous with thallus. Proper exciple free, (moderately) 

thin, hyaline to pale yellowish internally, yellowish to brownish or grayish marginally, nonamyloid. 

Hymenium up to c. 200 µm high, non-inspersed, moderately conglutinated, 

paraphyses parallel to slightly interwoven, unbranched, tips slightly thickened, lateral 

paraphyses present, usually conspicuous, up to c. 40 µm long, columellar structures absent. 

Epihymenium moderately thin, hyaline, with grayish to brownish granules and small crystals. 

Asci 4-8-spored, tholus (moderately) thick, usually not visible at maturity. Ascospores 

typical, (moderately) large, submuriform in younger stages to eumuriform at maturity, cell 

walls thick to very thick, endospore usually only slightly thickened, with thin to moderately 

thick, often distinctly irregular halo, hyaline, non-amyloid to faintly amyloid, predominantly 

fusiform to ellipsoid to more rarely clavate, ends narrowed-roundish to subacute, loci 

predominantly roundish, subglobose to flattened-subglobose to more rarely lentiform, end 

cells usually subglobose to slightly conical, septae thin to moderately thick, distinct


throughout development, ±regular, 60-140 x 15-30 µm with 12-22 x 1-6 loci. Pycnidia not 

seen (reported by Frisch 2006, for descriptions see there). 



lepadinum grows on tree bark in rainforests, wet 

and dry sclerophyll forests, heath shrubs, Karriforests 

and swamplands in temperate to subtropical 

regions in altitudes ranging from sea level 

to 1500 m. It is the most frequently collected 

species in southern Australia and widespread in 

temperate, oceanic Australia. It is the only species 

of the family with trentepohlioid photobiont 

occurring in south-western Australia. It further 

occurs in southern Queensland, New South Wales 

(incl. Lord Howe Island), Victoria and Tasmania. It 

is a subcosmopolitan species that is known from all 

continents except Antarctica (Frisch, 2006). 

NOTES – This species is characterized by a thin 

thallus, distinctly emergent, large ascomata with 

free, basally yellowish proper exciple and an entire, 

roundish thalline rim margin, (moderately) large, 

indistinctly muriform, hyaline, non- to faintly 


T. lepadinum. 

amyloid, thick-walled ascospores and the absence of secondary compounds. Several species 

of the Thelotrema s. str. group are morphologically close, particularly T. crassisporum (see 

there for differences). The temperate taxa T. subtile and T. suecicum are also similar (the latter 

often found growing in immediate proximity), both, however, are distinguished by smaller 

ascomata and transversely septate ascospores. Thelotrema adjectum has similar ascospores 

but differs in its morphology, T. lepadodes differs in ascospores that turn brownish at 

maturity (see also under that species). Thelotrema subgranulosum from Tasmania might 

represent an additional synonym, unfortunately the type collection was not available and is 

probably lost, see also under this species (appendices, part 5. 2.). 

SPECIMENS EXAMINED – Australia, Western Australia: The Cascades: 8 km S of Pemberton, Elix 10703 

(CANB); 4 km S of Pemberton, Elix 41100, 41101, 41112 (CANB), Lumbsch, Elix & Streimann 10789 a, 10791 

c (F). Warren NP.: 10-11 km SW of Pemberton, Elix 41233, 41238 (CANB); At Warren River, Tibell 14027 

(UPS). Near jct. of Heartbreak Trail and Old Vasse Rd., 10 km WSW of Pemberton, Kantvilas & Jarman 349/92 

(HO, PERTH). Darling: Karri forest, Pemberton, Baird 19741 (MEL); Beedelup Falls, WNW of Pemberton, 

Beauglehole 1035131 (MEL). Queensland: Mt. Mee SF., about 1 km NW of the forest station, Hafellner 16869 

(GZU). Gambubal SF., Cons Plain (Bald Mtn.) E of Emu Vale, Hafellner 16399, 16402 (GZU). New South 

Wales: Mt.Warning, track to and on summit, Mangold 18 a, l (F). Border Ranges NP., NE of Wiangaree: 

McPherson Range, uppermost part of Gradys Creek, Hafellner 16653 (GZU); Tweed Range, The Pinnacle, 

Hafellner 19180 (GZU). Nightcap NP., Mt. Nardi/Mt. Matheson Track, Mangold 22 j (F). Dorrigo SF., near 

Dorrigo, Hale s.n. (US). Dorrigo NP., Never Never Picnic Area and Rosewood Creek Track, Mangold 24 k, q 

(F). Gully W of radar tower adjacent to New England NP., Rogers 10312 (BRI). New England NP.: Above 

visitors' center, Hale 58774, 58778, 58782, 58834 (US); Track to Wrights Lookout, Elix 33931 (CANB); Radar 

Station Site, 1,5 km NW of Point Look Out, Kalb & Wiliams 20397, 20398, 20428, 20435, 20446, 20449, 

20451, 21213, 21232, 21397, 21927 (hb. Kalb). Styx River SF., 85 km E of Armidale: Kalb & Wiliams 21674, 

21676 (hb. Kalb); Softwood Rd., Rogers 10059 (BRI). C. 1 km W of Mt. Banda Banda, Kantvilas s.n. (NSW- 

219642). Barrington Tops SF., 40 km WNW of Gloucester: Dilgry River, Dilgry Circle Rd., Elix 24939, 24968 

(CANB); Streimann 44614 (B, CANB); Barrington Tops Forest Rd., Elix 24870, 24875 (CANB). Barrington 

Tops NP.: Gloucester Tops: Kantvilas s.n. (NSW-226538), W of Darby Munro Hut, Filson 1035799 (MEL); 

Thunderbolt Lookout,. (NSW-231139); Mt. William, Kantvilas 287/88 (NSW-221286), (HO). Allyn River


Forest Park: Start of track to Burraga swamp, Kantvilas s.n. (NSW-216962); Burraga Swamp, Kantvilas 165/88 

(HO, NSW-219642), 229/88 (HO). Old mill site c. 6 km N of Simsville and c. 30 km NW of Stroud, Filson 

1048795 (MEL). Duck Creek Rd., 22 km WNW of Buladelah, Elix 24412, 22414 (CANB). Trail along bank of 

Mill Creek, 50 km NW of Sydney, K. & A. Kalb 34271 (hb. Kalb). Blue Mountains NP.: Mt. Wilson: Weber 

220972, 229409 (COLO), Ewers 3095 (CANB), Kantvilas s.n. (HO), Chimney Cottage, K. & A. Kalb 20485, 

20578 (hb. Kalb); Near Blackheath, below Bridal Veil Falls, Hale 58671, 58675 (US); Cathedral Reserve circuit 

walk, Stajsic 2147425 (MEL). Mt. Tomah c. 20 km N of Katoomba, Thor 6171 (S). Katoomba, track between 

Leura Falls and Giant Stairways, K. & A. Kalb 20580, 20582, 20587, 20588, 20598 (hb. Kalb). 'National Park' 

[Royal NP.], Cheel 539350 (NSW). Stanwell Park [S of Royal NP.], Cheel 539351 (NSW). Monga SF./NP.: 

Clyde Mt., below the rd., Elix 972 (CANB); Milo Road, Streimann 43597 (CANB); 27 km SE of Braidwood, 

Mangold 11 a, l, y, z (F); 4-6 km S of Monga along Mungarlow River, Miller 426, Verdon 1349, Elix 11714, 

22692, 22696, 30223 (CANB), Kalb & Elix 21811, 21817, 21852 (hb. Kalb), Mangold 13 e (F). Glenbog SF., 13 

km ESE of Nimmitabel: Headwaters, Bemboka River, Bemboka River Rd., Streimann 43881 (B, CANB); 

Fastigata Rd., Streimann 43849 (CANB). Rutherford Creek, 11-17 km SE of Nimmitabel, Streimann 16755, Elix 

24290, 40813, 40840 (CANB). Brown Mt., 30 km SE of Nimmitabel, Kalb & Elix 30427, 30430, 30437, 30443 

(hb. Kalb). Lord Howe Island, Track to Goat House Cave, Elix 42187 (CANB). Victoria: Uncertain localities: 

'Waterloo Gypsland' [ =Waterloo Bay, Wilsons Promontory], Paton (BM-761914, -761915); 'Cunningham', 

Wilson s.n. (NSW-603860). Croajingolong NP., Double Creek Nature Walk, Mangold 9 b (F). Result Creek, 

Bendoc-Orbost Rd., 13 km SW of Bendoc, Elix 19837, 19839, 19871, 19873, 24146, Streimann 36530 (CANB). 

Hensleigh Rd., Queensborough River, 13 km SE of Bendoc, Elix 24081 (CANB). From Headwaters of West 

Errinundra River, Errinundra Plateau, Chesterfield 1060710, 1060741, 1060720, 1060178 (MEL). Errinundra 

NP.: Goonmirk Rocks, 14 km S of Bendoc, Elix 21877, 24190, 24204 (CANB); Coast Range Rd., 18 km SSE of 

Bendoc, Elix 21929, 24027 (CANB); Hammonds Rd., 19 km SSW of Bendoc, Elix 24169 (CANB). Jct. of Coast 

Range Rd. and Cobon Rd., 18.5 km SSE of Bendoc, Streimann 36708 (CANB). Ellery Camp, 30 km SSW of 

Bendoc, Elix 21442 (CANB). Club Terrace-Combienbar rd., 6 km N of Club Terrace, Elix 19516 (CANB). 

Bemm River, Princes Hwy., 8 km SSW of Club Terrace, Elix 19465 (CANB). Fernshaw at Watts River, NE of 

Healesville, Hafellner 18397 (GZU). Bonang rd., NE of Orbost, Mangold 8 a, f, h (F). Arte River, 30 km NE of 

Orbost, Elix 24212, 24215, 24223 (CANB). Lake Wat Wat, Dec. 1888, Wilson s.n. (NSW-518306, -539317). 

Lake Tyers, Mar. 1888, Wilson s.n. (NSW-539319). Lakes Entrance, Wilson s.n. (NSW-539328). Baw Baw NP., 

Mt. Erica carpark at start of walking track to Mt. Erica summit, Ford 2075897 (MEL). Moe Swamps [Moe], s.c. 

(MEL-26230). Strzelecki SF.: N of Grand Ridge Rd., 24.5 km SSE of Taralgon, Elix 29772 (CANB); Grand 

Ridge Rd., c. 3 km E of Gunyah Junction, 17 km NE of Foster Elix 29824 (CANB). Head of Franklin River, 

Thora-Gunyah Rd., 16 km NE of Foster, Streimann 65294 (B, CANB). [Tarra-]Bulga NP., Lyrebird-Ash Track, 

26 km ESE of Traralgon, Elix 29742, 29765 Streimann 51644 (CANB). Korumburra, 26. Jan. 1894, Wilson s.n. 

(NSW-539330). Wilsons Promontory NP.: Chinamans Creek, McVean 6557 (COLO); On Sealers Cove hiking 

track, Mangold 6 a, c, e, i, m (F). 90 km ENE of Melbourne, Cumberland Falls scenic reserve, Cora Lynn Falls, 

Thor 6030 (S). Wallaby Ck. Reserve, nr. Kinglake West, Willis 1023625 (MEL). Cumberland Valley at falls, c. 

16 km E of Marysville, Filson & Mayrhofer 508 (BM, CANB, H, US). Headwaters of Whitehouse Creek 

between Keppels Hut and Mt. Margaret, Filson 1023602 (MEL). Mt. Margaret Saddle, Blue Range, Filson 

1052096 (MEL). Along rd. to Lake Mountain [nr. Yarra Ranges NP.], Kantvilas s.n. (HO). Cement Creek Scenic 

Reserve, 5.5 km N of Warburton, Lumbsch, Dickhaeuser & Streimann 10974 a (F). River Watts [Yarra Ranges], 

1891, Wilson s.n. (NSW-539318). Black Spur [Yarra Ranges], Wilson s.n. (NSW-170675), 20. Mar. 1885 

(NSW-539321), 23. Mar. 1885 (NSW-539315), Feb. 1888 (NSW-603849, -603850), 1888 (NSW-603841, - 

603857, -603865), Jan. 1890 (S), 12. Nov. 1900, Bastow s.n. (MEL-26232). Warburton, Wilson s.n. (NSW- 

603853), 28. Feb. 1902, Bastow s.n. (MEL-26168, -16233). Gembrook [W of Bunyip State Park], 27. Oct. 1901, 

Bastow s.n. (MEL-26234). Ferntree Gully [W of Melbourne]: 1875, Paton s.n. (BM-761910, -761913, -761923), 

28. Jan. 1892, Wilson s.n. (NSW-603859), Bastow 26166, 26231, 724547 (MEL). Ottway Range: Ewers 7260 

(CANB); Todds Corner, Mt. Sabine Rd., Elix 25862 (CANB); Arkins Creek, watercatchment area in the Otway 

District, Filson 1023605 (MEL); gray River Rd., Angahook-Lorne SF., 17-28 km NE of Apollo Bay, Streimann 

58867, 58945 (CANB), 58962 (B, CANB); 14 km N of Apollo Bay, Mangold 5 h (F); Beauchamp Reserve, 

Otway SF., 13 km NNW Apollo Bay, Streimann 58839 (CANB); Skene Creek-Colac rd., 13 km ENE of Apollo 

Bay, Streimann 42860 (CANB); 10 km N of Apollo Bay, Mangold 3 b, f, I (F); Barham R. Rd.-Apollo Bay, 

Ewers 639920 (CANB); Binns Rd., 8 km WNW of Apollo Bay, Streimann 42672 (CANB); Maits Rest, Ocean 

Rd., 10 km W of Apollo Bay, Streimann 58645 (CANB). Ottway NP.: 10 km W of Apollo Bay, Mangold 1 a 

(F); Sandy Ridge Track Spur, 12 km W of Apollo Bay, Elix 25920, 25929, 25935 (CANB); Lighthouse Rd., 11 

km WSW of Apollo Bay, Curnow 1344 (CANB). Johanna River, Melba State Park, Lavers Hill, 26 km WNW of 

Apollo Bay, Streimann 58485, 58509 (CANB). Melba Gully, Ewers 3436, 3437 (CANB). Cobden, Devils Gully, 

20. Jan. 1885, Wilson s.n. (NSW-539316). Lake Elingamite [SW of Cobden], 15. Jan. 1885, Wilson s.n. (NSW- 

539320). Tasmania: Furneaux Group, Strezelecki Peaks, Flinders Island, Bass Strait, Filson 1023573 (MEL). 

Mt. Victoria Track, Kantvilas 54/81 (BM, HO). South Sister, near summit, Kantvilas 392/04, 409/04 (HO). 

Betw. Pyengana and Weldborough, Ewers 931 (CANB). Mt. Arthur, 23. Feb. 1891, Wilson s.n. (BM-726585, 

NSW-539394, UPS). Cheshunt [State Forest], 1867, Archer s.n. (BM-761894, -761896, -761897, -761898, -


761899, -761900, -761985). Near Projection Bluff, Lake Highway, Kantvilas 157/80 (BM, HO). Quamby Bluff, 

c. 15 km SSE Deloraine, along track from Lake Highway, Wedin 3135 (UPS). Warners Sugarloaf, 18.5 km S of 

Deloraine, Curnow 2023 (CANB). Gog Range, Mole Creek- Paradise Rd., 13 km SSE of Sheffield, Elix 26752, 

26757, 26762 (CANB). Rd. from Daisy Dell to Murchison Hwy., 17 km W of Daisy Dell, Elix 26804, 26833 

(CANB). Robbins Island Track, just N of Denium Hill, 25 km NW of Smithton, Elix 40257, 40262, 40270, 

40277 (CANB). Golden Valley, nr. Chestnut, Bratt 2428 (BRI). Sumac Spur 2, S of Arthur river, Kantvilas s.n. 

(BM-761920). Hellyer Gorge, McVean 6835 (CANB). Murchison Hwy., 57 km S of Bass Hwy., S of Hellyer 

Gorge, Hale 68793 (US). 6 km S of Warratah Hwy. jct. with the Murchinson Hwy., Hale 68727 (US). Doherty's 

Cradle Mountain Hotel grounds, Kantvilas 417/03 (HO). Cradle Mountain Lodge, 37 km NE of Rosebery, Elix 

23747 (CANB). Along the 'king billy track', Cradle Mountain Lodge, Elix 23598 (CANB). SE foothills of Mt. 

Black, Kantvilas 17/89 (HO). Near Williamsford, 6 km SSW of Rosebery, Elix 26874, 26880 (CANB). Anthony 

Rd., Kantvilas 194/91 (M), 577/92 (HO). Cradle Mtn. Lake St Clair NP., Pelion Plains, 1 km SW of Pelion Hut, 

Kantvilas 211/92 (HO). Corinna, Kantvilas 55/82 (HO). Zeehan, Fitzgerald 539325 (NSW). 10 km W of 

Strahan, along the Queenstown rd., Elix 5647 (CANB). 3 km S of Teepookana, Kantvilas 665/90, 670/90 (HO). 

Timbs Track, 27 km WSW of Maydena, Elix 27142, 27145 (CANB). Scotts Peak Rd., Kantvilas 433/80 (HO). 

W of Tahune Bridge, "Strips" Coupe CFI Plot in the Warra SST, Kantvilas s.n. (HO). Along Cracroft River near 

Cracroft Hut (Cracroft Crossing), Filson 1023608 (MEL). Mount Field NP.: Near Russell Falls, 55 km WNW of 

Hobart, A. & M. Aptroot 23297 (ABL); Lyrebird Trail, Hale 68758 (US). Myrtle Forest, Bratt s.n. (BM 761917). 

Lonnavale, Bratt s.n. (BM 761916). Track to Kermandie Falls, Gray s.n. (HO). SW Deadmans Bay, South 

Coast, Moscal 14095 (HO). De Witt Island, Elix 3332, 3340 (CANB). Five Road, Florentine Valley, Kantvilas 

289/80 (B, HO). Mt. Wellington: Wilson s.n. (NSW-539322); East slope, A. & M. Aptroot 22806 (ABL). 

Wielangta Rd., site E12 [WNW of Hobart], Kantvilas s.n. (HO). Uncertain locality: Turner s.n. (BM-761902). 

'Shirley Book', p. 121 (BRI-AQ721649, -AQ721650, -AQ721651). [Herb. Kremp., 1883], Hochstetter s.n. (M- 

34706). 

Thelotrema lepadodes Tuck. 

Proc. Amer. Acad. Arts Sci. . 5: 405 (1862). Leptotrema lepadodes (Tuck.) Zahlbr., Cat. Lich. Univ. II: 636 

(1923). Type: Cuba, Filanthropia, Wright s.n. (FH-Tuck.-lectotype, selected by Hale [1978b: 52]; FH-Tuck.isolectotype). 

Thelotrema monosporum var. patulum Nyl., Acta Soc. Sci. Fenn. 7: 452 (1863a). Leptotrema patulum (Nyl.) 

Müll. Arg., Bull. Herb. Boissier 3: 315 (1895b). Type: Colombia (Nova Granata), 1860, Lindig s.n. (H- 

Nyl.22715-lectotype, selected by Hale [1978b: 52]; H-Nyl.22717-isolectotype). 

Thelotrema disciforme Leight., Trans. Linn. Soc. London 27: 170 (1869). Phaeotrema disciforme (Leight.) 

Hale, Smithsonian Contrib. Bot. 16: 29 (1974b). Type: Sri Lanka (Ceylon), Central Prov., Thwaites C.L. 128 

pr.p. (BM!-lectotype, selected by Frisch [2006: 300]; H-Nyl. 3851-isolectotype). 

Leptotrema aemulum Müll. Arg., Bull. Herb. Boissier 3: 316 (1895). Type: Australia, Queensland, 1887, 

Knight 27 + 295 [same specimen] (G!-[Knight 295]-lectotype, selected by Hale [1972 in herb.]; G!-[Knight 27]isolectotype). 

Leptotrema bisporum Szatala, Ann. Mus. Nat. Hungar. n.s.7: 30 (1956). Type: Solomon Islands, ('Nova 

Guinea'), Berlinhafen, Seleo Isl., Biro 254 (BP!-holotype). 

[For additional synonymy see Frisch 2006 (as T. monosporum), but see also note on T. saxatile below.] 


Thallus epi- to predominantly hypophloedal, very thin to thin, up to c. 150 µm high, pale 

gray to yellowish-gray. Surface dull to somewhat glittering, smooth to more often roughened, 

often due to protuberant substrate, predominantly continuous, predominantly unfissured, 

sometimes appearing fissured due to substrate structure. Cortex structures absent or covered 

by a thin, incontinuous protocortex up to c. 15 µm thick. Algal layer poorly developed, 

continuous to incontinuous, calcium oxalate abundant to more rarely sparse, usually small to 

more rarely large, scattered to more rarely clustered. Vegetative propagules not seen. 

Ascomata conspicuous to inconspicuous, (moderately) large, up to c. 900 µm in diam., 

roundish, apothecioid, ±sessile, solitary to marginally fused, becoming ±distinctly emergent, 

hemispherical to urceolate with same surface than thallus. Disc becoming partly visible from 

surface, pale grayish to dark gray, distinctly pruinose. Pores moderately small to usually wide 

or gaping, up to c. 700 µm, roundish to more rarely roundish-irregular, entire to slightly split,


Fig. 138. Thelotrema lepadodes: growth habit (A, B), ascomata (C, D), ascospores (E), younger 

ascospore (F) and mature ascospore (G). A.: G-lectotype of L. aemulum; B., E.-G.: Lumbsch & 

Guderley 11138 d; C.: BM-lectotype of T. disciforme; D.: BP-holotype of L. bisporum. Bar= A: 1 

mm; B, C: 0.8 mm; D: 0.4 mm; E: 20 µm; F, G: 9 µm. 

proper exciple apically to entirely visible from surface, off-white, sometimes shrunken, 

incurved to more rarely erect. Thalline rim margin roundish, wide to gaping, entire to 

somewhat split or eroded, thin to moderately thick, incurved to rarely slightly erect, thalline 

rim concolorous with thallus to sometimes darker than thallus. Proper exciple becoming free, 

(moderately) thin, predominantly hyaline internally, pale to dark brown marginally,


sometimes amyloid at the base. Hymenium up to c. 200 µm high, non-inspersed, distinctly 

conglutinated, paraphyses parallel or slightly interwoven, unbranched, tips unthickened to 

slightly thickened, lateral paraphyses present, inconspicuous, up to c. 20 µm long, columellar 

structures absent. Epihymenium (moderately) thick, hyaline, with fine to coarse grayish 

granules and small crystals. Asci 2-8-spored, tholus (moderately) thick, thin when mature. 

Ascospores moderately large, (sub-)muriform, cell walls moderately thin in younger stages to 

thick at maturity, endospore (moderately) thick, covered by a thin, smooth to somewhat 

irregular halo, hyaline to slightly brown at early maturity, becoming distinctly brown, nonamyloid 

to faintly to more rarely moderately amyloid, rarely cylindrical to usually fusi- or 

claviform, with roundish to subacute ends, loci roundish, subglobose to lentiform or oblong, 

end cells hemispherical to conical, transverse septae moderately thin to thick, distinct, regular, 

40-100 x 10-25(30) µm with c. 12-20(22) x 1-5(6) loci. Pycnidia not seen. 



lepadodes grows on tree bark in coastal rainforests 

and mangroves in altitudes ranging from sea level 

to 20 m. It is moderately common occurring in 

northern to north-central Queensland. The pantropical 

species is also known from Polynesia, 

Hawaii, the Neotropics, Africa, Sri Lanka, 

Philippines and Solomon Islands (Frisch, 2006, as 

T. monosporum). 


predominantly hypophloedal thallus, open-pored, 

±emergent ascomata, moderately large, (sub-) 

muriform, thick-walled, indistinctly brownish, nonto 

faintly amyloid, fusi- to claviform ascospores in 

8-spored asci, and the absence of secondary 

compounds. Similar species include T. lepadinum 

and T. conveniens, which can be distinguished by 

constantly hyaline ascospores (T. lepadinum), and 

monosporic asci with larger (up to 200 µm long), 


T. lepadodes. 

thin-walled, distinctly amyloid ascospores (T. conveniens), respectively. Unfortunately the 

type of T. lepadodes was not available for study, however, it is referred to Frisch's (2006) 

detailed description of the species under the name 'T. monosporum' which agrees well with 

the Australian specimens. Frisch (2006) noted that since the H-Nyl. type of T. monosporum is 

in bad condition, the correct name for this species remains uncertain. Further he wrote (2006: 

303): "If it (…) represents a species separate from the collections (…) presently identified 

with the name T. monosporum, T. lepadodes is the oldest name available for these collections 

at species level." This approach is also supported by the fact that Frisch lists four tentatively 

included specimens that differ from his concept of T. monosporum (=T. lepadodes) by single 

spored asci and elliptical ascospores. It is assumed that the specimens described belong to T. 

saxatile since the anomalous characters described agree well with the differences of T. 

lepadodes and T. saxatile. Frisch (ibid.) lists T. saxatile as a synonym to T. monosporum 

without having seen the type, referring to Salisbury (1972). However, it is here regarded as a 

distinct species. For a detailed list of differences between T. lepadodes and T. monosporum/T. 

saxatile see under these taxa. In Müller’s original description of L. aemulum (Müller 1895) he


noted a collection from Victoria, the sample in G (Knight 303), however, lacks any locality 

data. Since the species is tropical, the Victorian locality is most probably erroneous. 

SPECIMENS EXAMINED – Australia, Queensland: Thursday Island (N of York Peninsula), 1887, Knight s.n. 

(G-10194/5). Iron Range NP., c. 3.5 - 31 km from western boundary on track to Portland Rds., Hale 830010, 

830020 (US). Cape Tribulation Area: Myall Beach, Lumbsch & Mangold 1958 k, 19162 o (F); Track to Cape 

Tribulation Beach, Mangold 32 q (F). Cooga Beach, 5 km E of Mossman, Lumbsch & Mangold 19169 d (F). 

Cairns Airport, Ewers 8724 (CANB). Russell River NP., 8 km NE of Babinda, mangrove at Russell River, 

Lumbsch & Guderley 11138 d (F). 3 km S of Forrest Beach, 16 km SE of Ingham, Elix & Streimann 15924, 

15933 (CANB). Three Mile Creek, 5 km N of Townsville, Elix & Streimann 20049 (CANB). Mangrove swamp 

on rd., just before Shute Harbour, Hale 831576 (US). Conway Range NP., near Shute Harbour-Airlie Beach, 

Hale 832127 (US). Cape Hillsborough NP., NW of Mackay, Hale 831494 (US). Mimeo Swamp, Sunset Beach 

(N of Mackay), Stevens 2545 (BRI). Notch Point rd., S of Cape Palmerston NP., Hale 832107, 832124 (US). 

Uncertain locality, Knight 303 (G). 

Thelotrema leucophthalmum Nyl. 

Bull. Soc. Linn. Normand. 2(2): 39 (1868). Type: New Caledonia, Lifu, Loyalty Isl., 1864, Thiebaut s.n. (H- 

Nyl. 22518!-lectotype, selected by Hale [1972 in herb.]). 

Leptotrema albocoronata C. Knight in Shirley, Proc. Roy. Soc. Queensland 6: 192 (1889). Type: Australia, 

Queensland, Sankey's Shrub (Brisbane), Shirley 509 (WELT!-lectotype, selected here; BRI-‘Shirley Book’, p. 

23, n. 8 [BRI-AQ721248]!-isolectotype). 


Thallus epi- to hypophloedal, thin to moderately thick, up to c. 300 µm high, pale 

yellowish-green to yellowish-brown to pale olive. Surface waxy, smooth, continuous, 

unfissured. True cortex present, yellowish, up to c. 80 µm thick, consisting of irregular to 

periclinal hyphae. Algal layer continuous to incontinuous, well developed, calcium oxalate 

crystals abundant, variable in size, often clustered. Vegetative propagules not seen. Ascomata 

very conspicuous, (very) large, up to c. 2 mm in diam., roundish, slightly irregular in fused 

ascomata, apothecioid, erumpent, solitary to fused, moderately emergent, conical to broad 

annular. Disc partly to rarely entirely visible from surface, pale grayish to pale flesh-colored, 

coarsely pruinose. Pores becoming wide to gaping, up to c. 1.5 mm in diam., roundish to 

moderately irregular, inner and apical proper exciple becoming visible from surface, fused to 

free, ± jagged, whitish, incurved to recurved, somewhat shrunken only in entirely free proper 

exciples. Thalline rim margin predominantly thick, roundish to moderately irregular, split to 

lacerate, ±coarsely lobed, sometimes indistinctly layered, becoming ±distinctly eroded, 

whitish, pruinose, thalline rim concolorous and with same surface as thallus, predominantly 

erect to recurved. Proper exciple moderately thick, hyaline internally, (pale) yellowish-gray to 

brownish marginally, often with calcium oxalate crystal inclusions, apically usually covered 

by grayish granules, non-amyloid to rarely faintly amyloid at the base. Hymenium up to c. 

130 µm high, non-inspersed, moderately conglutinated, paraphyses ±bent, ±interwoven, 

unbranched, tips slightly thickened and somewhat irregular, lateral paraphyses present, 

usually very inconspicuous, up to c. 25 µm long, true columella absent, in fused ascomata 

sometimes columella-like structures present. Epihymenium ±thick, hyaline to pale grayishbrown, 

with grayish granules and sometimes small crystals. Asci 6- to 8-spored, tholus 

moderately thick, thin when mature. Ascospores moderately small, distinctly muriform in 

late stages, cell walls (moderately) thick, endospore moderately thick, sometimes with a 

moderately thin, irregular halo, hyaline, non-amyloid to rarely faintly amyloid, oblong to 

ellipsoid to more rarely fusi- or claviform, with roundish to narrowed-roundish ends, loci 

roundish to slightly angular, subglobose to oblong or irregular, transverse septae moderately 

thin, distinct, regular, 30-50(60) x 10-15 µm with 8-16 x 1-5(6) loci. Pycnidia not seen.


Fig. 140. Thelotrema leucophthalmum: growth habit (A, B), ascomata (C), ascoma section (D), 

immature ascospores (E) and mature ascospore (F ). A., C., E.: WELT-lectotype of L. 

albocoronata; B., D., F.: Tibell 14660. Bar= A: 1.7 mm; B: 1.3 mm; C: 1 mm; D: 120 µm; E: 10 

µm; F: 6 µm. 


constictic (majors), hypostictic (major to trace), α-acetylconstictic and cryptostictic (traces) 

acids. 

ECOLOGY AND DISTRIBUTION – Thelotrema leucophthalmum was collected in Australia 

growing on tree bark in (sub)tropical rainforests and wet sclerophyll forests in altitudes 

ranging from sea level to 900 m. It is moderately common and wide-spread in northern to 

southern Queensland. It is also known from New Caledonia and the Andman Islands (Sethy & 

al., 1987) and newly reported here for the Neotropics (Puerto Rico), being pantropical. 

NOTES – The species is characterized by a dark, waxy, corticate thallus, conspicuous, large 

ascomata with a whitish, split to lacerate and ±eroded, pruinose thalline rim margin, 

moderately large, indistinctly muriform, hyaline, non-amyloid to faintly amyloid ascospores


and the stictic acid chemosydrome. It is similar to 

T. cupulare and T. thesaurum, both readily distinguished 

by smaller and larger ascospores 

respectively (up to up to 30 µm in T. cupulare, up 

to 130 µm in T. thesaurum). 


Tribulation Area: Teahouse at intersection of Buchanan Creek 

Rd. and Cape Tribulation Rd., Hale 831180 (US); Cape 

Kimberley, Lumbsch & Mangold 19164 q (F). Julatten, Ewers 

8446 (CANB). Stallion Pocket logging area, 14 km from 

Gillies Hwy and 1 km E from Mulgrave River Forestry rd., S 

of Gordonvale, Hale 832326 (US). Mt. Hypipamee NP., 

Dinner Creek Falls, S of Atherton, Hale 831369 (US). 

Conway Range NP.: Near Shute Harbour-Airlie Beach, Hale 

71182 (US); At the campground 0.5 km E of Shute Harbour, 

Tibell 14660 (UPS). Conway SF., 18 km E of Proserpine, Elix 

& Streimann 20210 (CANB). Cape Hillsborough NP., NW of 

Mackay, Hale 831290 (US). Kalpower Forest Drive, 40 km 

NE of Monto, SW of Gladstone, Hale 831500, 831649 (US). 

Wooroi SFP., W of Teewantin, Hale 832168 (US). Blackall 

Ranges, Wilson s.n. (NSW-539400). Brisbane, Bailey 143 

(BM, BRI), Bailey s.n. (BM -726590). Porto Rico, Indiera Fria, near Maricao, Britton, Cowell & Brown 4501 

(US). 

Thelotrema monosporum Nyl. 

Ann. Sci. Nat., Bot., 4(15): 46 (1861b). Type: New Caledonia, Pancher s.n. (H-Nyl.22709!-lectotype, 

selected by [Hale 1981: 260]). 



T. leucophthalmum. 

Thallus predominantly hypophloedal, very thin to thin, up to c. 100 µm high, pale 

yellowish-gray to pale grayish-green. Surface dull, usually roughened, continuous, unfissured. 

Cortex structures absent or covered by an incontinuous protocortex up to 15 µm thick. Algal 

layer moderately well developed, continuous to incontinuous, calcium oxalate usually 

abundant, sometimes sparse, small to large, scattered or clustered. Vegetative propagules not 

seen. Ascomata inconspicuous, moderately small to moderately large, up to 600 µm in diam., 

roundish, somewhat peri- to predominantly apothecioid, ±sessile, solitary to marginally fused, 

becoming ±distinctly emergent, predominantly hemispherical. Disc not visible from surface to 

very rarely becoming somewhat visible, usually only in defective ascomata, grayish, 

moderately pruinose. Pores (moderately) small, up to c. 300 µm, roundish to more often 

roundish-irregular, proper exciple apically visible from surface, entirely free, off-white to 

whitish, often shrunken, predominantly incurved to more rarely somewhat erect. Thalline rim 

margin roundish to irregular, predominantly moderately small, entire to slightly split, 

moderately thin to thick, incurved, concolorous with thallus to more rarely brighter than 

thallus. Proper exciple ±free, moderately thin to moderately thick, hyaline to pale yellowish 

internally, yellowish-brown marginally, apically often brownish to dark-brown or dark-gray, 

sometimes amyloid at the base. Hymenium up to c. 200 µm high, non-inspersed, moderately 

conglutinated, paraphyses parallel or slightly interwoven, unbranched, tips moderately to 

distinctly thickened, lateral paraphyses present, often inconspicuous, up to c. 20 µm long, 

columellar structures absent. Epihymenium variably thick, hyaline, usually with small crystals 

and fine to coarse grayish granules. Asci 1 to 4-spored, tholus (moderately) thin, not visible at 

maturity. Ascospores moderately large, eumuriform, cell walls and endospore predominantly


(moderately) thin, in younger stages often distinctly thickened parts present, non-halonate, 

brown at early maturity, non-amyloid to very faintly amyloid in young ascospores, ellipsoid 

to predominantly (broad-)fusiform, very rarely somewhat claviform, with narrowed-roundish 

ends, loci predominantly roundish to slightly angular, subglobular to somewhat oblong, 

transverse septae thin, distinct, regular, 40-90(100) x 15-30 µm with c. 14-26 x 2-8 loci. 


Fig. 142. Thelotrema monosporum: growth habit (A), ascomata (B), immature, mature and overmature 

ascospores (C, D) and ascospore detail (E). A., D., E.: H-lectotype; B., C.: Mangold 32 u. 

Bar= A: 0.75 mm; B: 0.6 mm; C: 30 µm; D: 15 µm; E: 10 µm. 




monosporum occurs in tropical rainforests in low 

altitudes ranging from sea level to 20 m on tree 

bark. It is rare in Australia being restricted to 

northern Queensland. The distribution is unclear, 

given the taxonomic difficulties (discussed below). 

The type was collected in New Caledonia. 


mainly hypophloedal and ecorticate thallus, 

moderately large, muriform, brown, thin-walled, 

non- or faintly amyloid ascospores in 1-4-spored 

asci and the absence of secondary metabolites. 

The circumscription of this species has differed 

widely between authors [see Frisch, 2006; 


T. monosporum.


Matsumoto, 2000; Purvis & al., 1995 (T. macrosporum); Harris, 1990 (T. monospermum)]. 

The main reason for the problems lie in the poor conidition of the type of T. monosporum and 

consequently, different concepts of its differences to T. lepadodes and T. saxatile. While some 

authors consider these as synonymous with T. monosporum, they are regarded as distinct 

species here (see also under that species). The Nylander-type specimen has a moderately thin, 

ecorticate, roughened thallus with numerous calcium oxalate crystal inclusions, small, slightly 

raised ascomata with small openings, a free proper exciple that is partly visible, several 

decayed, but also a few intact ascospores that were distinctly brown, densely eumuriform, 

non-amyloid, broadly fusiform with rounded ends, with (moderately) thin cell walls, thin 

endospore and septae, 70-90 x 20-30 µm in size (Fig. 143, D, E). The number of spores per 

ascus, structure of lateral paraphyses or exciple structure could not be studied due to the poor 

material. However, ascomata are present that agree with the protologue. Thelotrema 

lepadodes differs in having 4-8 spored asci, and more distinctly tapered ascospores with 

±subacute ends. Additionally, the ascospores of T. lepadodes become brown late in 

development and are often distinctly amyloid. Another problematic species is T. saxatile see 

under this species for differences. Galloway (1985) considers Ascidium attenuatum from New 

Zealand as a synonym of T. monosporum, which could not be checked since the type material 

is lost and only a drawing of a single ascospore (selected as the type by Galloway) is 

available. Ascidium attenuatum is considered a nomen dubium (see appendix). It is 

questionable that it really is conspecific with T. monosporum, since the drawing as well as the 

protologue state transversely septate ascospores. 


Portland Rds., Hale 832441 (US). Cape Tribulation Area: Myall Beach, Lumbsch & Mangold 19158 g, i, j, v, w, 

19160 d, 19161 n, x, 19162 i, p, Mangold 31q (F); Track to Cape Tribulation Beach, Mangold 32 u (F). 

Thelotrema myriocarpum Fée 

Essai Cryptog. Écorc. Officin.: 94 (1824). Myriotrema myriocarpum (Fée) Hale, Mycotaxon 11: 134 (1980). 

Type: 'America meridionali, ad corticem Cinchonae rubrae' (PC-lectotype, selected by Hale [1978:45]; G-Fée 

242-, L-Buse!-isolectotypes). 


Thallus epi- to slightly hypophloedal, moderately thick, up to c. 400 µm high, pale 

yellowish green to usually (pale) olive. Surface shiny, smooth, continuous to rugose and/or 

verruculose, unfissured. True cortex present, continuous, consisting of periclinal hyphae, up 

to 30 µm thick. Algal layer well developed, continuous, often becoming incontinuous due to 

crystals inclusions, calcium oxalate crystals very abundant, small to large, scattered or 

clustered, sometimes forming column-like structures. Vegetative propagules not seen. 

Ascomata variable, inconspicuous, (moderately) small, up to 350 µm in diam., roundish, perito 

apothecioid, solitary, immersed to slightly raised. Disc usually becoming partly visible 

from surface, pale grayish to pale flesh-colored, epruinose. Pores (moderately) small, up to c. 

250 µm in diam., roundish to sometimes slightly irregular, apical proper exciple becoming 

±visible from surface, predominantly fused to rarely partly to entirely free, entire to more 

rarely slightly split, ±whitish to pale brownish, incurved. Thalline rim margin (moderately) 

thin, entire, roundish, usually brighter than thallus to whitish, thalline rim incurved, 

concolorous with thallus to rarely grayish-brown, with same surface as (or not distinguishable 

from) main thallus. Proper exciple fused to partly free, predominantly thin, hyaline to pale 

yellowish internally to orange marginally, apically sometimes dark-brown, non-amyloid to 

weakly amyloid at the base. Hymenium up to c. 130 µm high, non-inspersed, distinctly 

conglutinated, paraphyses ±straight, ±interwoven, unbranched, tips thickened, lateral


paraphyses present, inconspicuous, up to c. 20 µm long, columellar structures absent. 

Epihymenium indistinct and thin, hyaline, rarely with grayish granules. Asci 8-spored, tholus 

(moderately) thick, moderately thin when mature. Ascospores small, (sub-)muriform, cell 

walls (moderately) thin, endospore moderately thick, non-halonate, hyaline, non-amyloid to 

rarely faintly amyloid (reddish), oblong to ellipsoid to somewhat claviform, with roundish to 

narrowed-roundish to rarely subacute ends, loci roundish to ±angular, in particular in younger 

stages, subglobose to oblong to predominantly ±irregular, transverse septae (moderately) thin, 

distinct, regular, 20-30(40) x 7-10 µm with 6-11(12) x 1-4 loci. Pycnidia not seen. 


stictic, (majors), α-acetylconstictic, cryptostictic, α-acetyl-hypoconstictic (minors to traces) 

and hypostictic (trace) acids. 

Fig. 144. Thelotrema myriocarpum: growth habit (A), ascomata (B), lateral paraphyses (C), and 

ascospores (D, E). A.-D.: Lumbsch & Mangold 19120 l; E.: Lumbsch & Mangold 19167 c. Bar= 

A: 1 mm; B: 0.7 mm; C: 20 µm; D, E: 5 µm. 

ECOLOGY AND DISTRIBUTION – Thelotrema myriocarpum grows on tree bark in tropical 

rainforests in altitudes ranging from sea level to 900 m. It is moderately common in northern 

Queensland. This is the first report for Australia and the paleotropics. The pantropical species 

was previously regarded as neotropical (Hale, 1978). 

NOTES – The species is characterized by a shiny, corticate, thick thallus, small ±immersed 

ascomata, indistinct lateral paraphyses, moderately small, hyaline, (sub-)muriform, 

predominantly non-amyloid ascospores with thin walls but thickened endospore and the


presence of the stictic acid chemosyndrome. The 

pantropical (but not known from Australia) 

'Thelotrema' glaucopallens is similar but differs in 

the absence of lateral paraphyses. A similar 

Australian species is T. cyphelloides, for 

differences see under this species. The Australian 

collections confirm well with the type except for 

smaller ascospores. In contrast to sizes of up to 40 

µm found in the type, the ascospores in the 

Australian material never exceed 30 µm in length. 

The measures found in literature range from up to 

50 µm (Salisbury, 1978) to 25 µm in Hale (1978), 

however, the latter author stated that he found 

length of up to 36 µm in the PC type. 


York Peninsula: Iron Range NP., 4-9 km from western 

boundary on track to Portland Roads, Hale 830022, 830044, 

830046, 830050, 830080, 832672, 832810 (US); McIlwraith 

Range, Rocky River, 60 km SE of Coen, Jones s.n. (MEL). 

Near Cedar Bay NP., on rd. to Cooktown, Mangold 34 k, p, s 

(F). Cape Tribulation Area: Cow Bay, Hale 831190, 832464 (US); Cape Kimberley, Lumbsch & Mangold 19167 

c, r (F). Mossman Gorge NP., Hale 831055, 832606 (US). Crystal Cascades, 5 km W of Cairns, Lumbsch & 

Mangold 19120 d, l (F). SW of Gordonvale, 25-30 km on Mulgrave River Forestry Rd., Hale 831015, 832266 

(US). Atherton Tablelands: Lake Euranoo, Lumbsch & Mangold 19127 g, j, l, w (F); Curtain Fig Tree SFP., Hale 

831220 (US); Along west boundary of Eacham NP., on rd from Atherton, Hale 831018 (US); Palmerston NP. 

(=south part of Wooroonooran NP.), Hale 831130 (US). Picnic area 2 km E of Cardstone on Tully River Rd. to 

Kareeya Power Station, Hale 830958 (US). Uncertain location: 'Shirley book', p. 23, n. 7 [as 'Leptotrema 

phaeosporum'], s.c. (BRI-AQ721247). 

Thelotrema nostalgicum Salisb. 

Lichenologist 5: 266 (1972). Type: Sri Lanka, Pidurutalga, 15.-22. Dec. 1879, Almquist s.n. (H-Nyl. 22741!lectotype, 

selected by Hale [1981: 260]; S!-isolectotype). 



T. myriocarpum. 

Thallus predominantly epi- to slightly hypophloedal, moderately, up to c. 200 µm high, 

(pale) greenish gray to (pale) olive or yellowish gray. Surface dull to slightly shiny, smooth to 

slightly rough, continuous to more often distinctly or strongly verruculose, unfissured to 

slightly cracked. Algal layer well developed, continuous, calcium oxalate crystals abundant, 

predominantly small, scattered or clustered. Vegetative propagules not seen. Ascomata 

±conspicuous, predominantly large, up to c. 1.3 mm in diam., roundish to irregular-roundish, 

sessile, peri- to apothecioid, solitary to marginally fused, becoming distinctly emergent, 

hemispherical to urceolate, with same surface as thallus. Disc not visible from surface to 

rarely partly visible, grayish, epruinose to slightly pruinose. Pores moderately small, up to c. 

300 µm in diam., roundish to slightly irregular, entire to somewhat split, apical proper exciple 

sometimes becoming visible from surface, fused to indistinctly free, rarely somewhat 

shrunken, whitish to off-white or slightly brownish apically, incurved. Thalline rim margin 

moderately thick to thick, becoming moderately wide, roundish to slightly irregular, 

predominantly entire to slightly split, incurved. Proper exciple becoming apically free, 

moderately thick to thick, hyaline to pale yellowish internally, (pale)brownish to yellowishbrown 

marginally, apically sometimes darkish-gray, sometimes ±distinctly amyloid at the


Fig. 146. Thelotrema nostalgicum: ascomata (A, B), immature ascospore (C), ascospore detail (D) and 

ascospore detail showing amyloid reaction (E). A.-E.: Hale 50356. Bar= A: 0.75 mm; B: 0.7 mm; 

C: 10 µm; D: 7 m; E: 12 µm. 

base. Hymenium up to c. 300 µm high, non-inspersed, distinctly conglutinated, paraphyses 

±parallel, unbranched, tips slightly thickened, lateral paraphyses present, conspicuous, up to 

c. 40 µm long, columellar structures absent. Epihymenium moderately thin to thin, hyaline, 

often with grayish granules. Asci 2-4-spored, tholus moderately thick to thick, thinning with 

maturity. Ascospores large to very large, transversely septate, cell walls thick to very thick, 

sometimes with a distinctly crenate to ±irregular outline, non-halonate, hyaline, 

predominantly faintly to moderately amyloid, cylindrical to fusiform or bifusiform, ends 

roundish to narrowed-roundish, loci regular, roundish, subglobose to lentiform or oblong, 

endcells hemispherical, septae moderately thin to moderately thick, regular, 80-240 x 10-25 

µm with 16-38 loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellowish-brown, C-, PD+ orange-red; containing fumarprotocetraric 

(major) and protocetraric (major to trace) acids. 

ECOLOGY AND DISTRIBUTION – Fide Kalb (2001), Thelotrema nostalgicum occurs on tree 

bark and in Australia is restricted to subtropical rainforests at altitudes ranging from 900 to 

1000 m. It is a very rare species occurring in the Queensland/New South Wales border region. 

This is the first report for Australia. Besides Australia, it is was reported from Sri Lanka 

indicating a palaeo(sub)tropical distribution. 

N OTES – This taxon is characterized by distinctly emergent ascomata with rather 

indistinctly free proper exciple, usually very large, transversely septate, hyaline, amyloid


ascospores with conspicuously thick cell walls and 

the presence of protocetraric acid chemosydrome 

compounds. It is very similar to T. nureliyum, but is 

distinguished by more distinctly emergent ascomata 

with less distinctly free proper exciple, slightly 

larger ascospores that occur by two to four per 

ascus (ascospores 4-8 per ascus with up to 220 x 25 

µm with up to 35 loci in T. nureliyum) and by the 

presence of protocetraric acid chemosydrome 

compounds (secondary compounds absent in T. 

nureliyum). In Australia the species is only known 

from two collections from the Lamington National 

Park (reported by Kalb, 2001), which, however, 

were not available for study. 

SPECIMENS EXAMINED – Sri Lanka, Nuwara Eliya, Hale 

50356 (US). 

Thelotrema nureliyum Hale 

Bull. Brit. Mus. nat. Hist. (Bot.) 8: 261 (1981). Type: Sri Lanka, Central Province, Hale 50292 (US-holotype, 

not seen). 



T. nostalgicum. 

Thallus variable, predominantly epi- to slightly hypophloedal, moderately thin to 

moderately thick, up to c. 300 µm high, (pale) greenish gray to (pale) olive or yellowish gray. 

Surface predominantly dull, smooth to roughened, sometimes porous, continuous to more 

often distinctly or strongly verruculose, sometimes forming subglobular, isidia-like structures, 

distinctly to strongly fissured. Thallus covered by a usually thin, ±continuous protocortex up 

to c. 25 µm thick. Algal layer well developed, continuous, calcium oxalate crystals abundant, 

predominantly small, scattered or clustered. Vegetative propagules not seen, isidia-like 

structures sometimes present (see above). Ascomata ±conspicuous, predominantly large, up to 

c. 1.2 mm in diam., roundish to irregular-roundish, apothecioid, somewhat erumpent, solitary 

to marginally fused, becoming moderately to distinctly emergent, hemispherical to urceolate, 

with same surface as thallus or sometimes more distinctly verruculose. Disc often becoming 

partly to rarely entirely visible from surface, pale grayish to whitish, ±strongly pruinose. 

Pores relatively small to moderately small, up to c. 400 µm in diam., roundish to slightly 

irregular, entire to split, apical to entire proper exciple becoming visible from surface, often 

somewhat shrunken, whitish to off-white or slightly brownish apically, concolorous to pale 

brownish towards the base, incurved to erect to more rarely slightly recurved. Thalline rim 

margin moderately thin to thick, becoming moderately wide to gaping, roundish to slightly 

irregular, predominantly entire to split, sometimes lacerate or eroded, often distinctly layered, 

incurved to erect, concolorous with thallus, in layered margins inner layer(s) usually more 

bright. Proper exciple becoming entirely free, (moderately) thick, hyaline to pale yellowish 

internally, (pale)brownish to yellowish-brown marginally, apically usually dark-gray and 

covered by granules, often distinctly amyloid at the base. Hymenium up to c. 250 µm high, 

non-inspersed, distinctly conglutinated, paraphyses ±parallel, unbranched, tips distinctly 

thickened, lateral paraphyses present, conspicuous, up to c. 40 µm long, columellar structures


absent. Epihymenium thick to very thick, hyaline, with grayish to dark gray granules and 

moderately small crystals. Asci 4-8-spored, tholus (moderately) thick, thin when mature. 

Ascospores variable, moderately large to very large, transversely septate, very rarely with a 

single longitudinal septum, cell walls thick to very thick, usually with a distinctly crenate to 

±irregular outline, non-halonate, hyaline, becoming faintly yellowish when decayed, 

predominantly faintly to moderately amyloid, cylindrical to fusiform or sometimes 

bifusiform, ends roundish to narrowed-roundish, loci regular, roundish, subglobose to 

lentiform or oblong to rarely biconcave, end cells hemispherical, septae moderately thin to 

moderately thick, regular, 60-220 x 10-20 µm with 12-35 (x 2) loci. Pycnidia not seen, fide 

Matsumoto & Deguchi (1999) present, immersed with dark pore area, conidia oblong, up to 

2.4 x 0.8 µm. 


Fig. 148. Thelotrema nureliyum: growth habit (A), ascomata (B), ascospores (C, D) and ascospore 

showing amyloid reaction (E). A., C., E.: Lumbsch & Mangold 19108 h; B., D.: Lumbsch & 

Mangold 19174 x. Bar= A: 2 mm; B: 1.5 mm; C: 15 µm; D: 10 µm; E: 12 µm.



nureliyum was collected in Australia on tree bark in 

(sub)tropical to warm-temperate rainforests and 

more rarely in wet sclerophyll forests in elevations 

ranging from 80 to 1500 m (mostly between c. 600 

and 1000 m). It is common and widespread in 

eastern Australia, occurring from northern Queensland 

to south-central New South Wales and Lord 

Howe Island. This is the first report for Australia. 

This paleotropical species has been recorded from 

India (Patwardhan & al., 1985), Sri Lanka and 

Japan (Matsumoto, 2000). 

NOTES – Thelotrema nureliyum belongs to the T. 

subtile-group and is characterized within this group 

by a thick, fissured and often strongly verruculose 

thallus, large ascomata with distinctly free proper 

exciple, and large, transversely septate, hyaline, 

±weakly amyloid ascospores that usually have a 

conspicuously thick cell wall. Thelotrema diplo- 


T. nureliyum. 

trema has similar ascospores, see under that species for differences. Among Thelotrema 

species with transversely septate ascospores only T. gallowayanum (ascospores brown with 

distinctly tapered ends, up to 400 µm long) has distinctly larger ascospores. Thelotrema 

nostalgicum has only slightly larger (up to 240 µm) ascospores but the asci are 2-4-spored 

(for further differences see under that species). 

SPECIMENS EXAMINED – Australia, Queensland: Thornton Range, CREB rd. (to Cooktown), about 5 km in 

from Daintree River crossing, NW of Mossman, Hale 831527 (US). Mt. Windsor logging area, E of old Forestry 

Camp, NW of Mossman, Hale 832329, 832474 (US). Mt. Lewis Rd. 10.5 km N from Kennedy Hwy., W of 

Mossman, Hale 830678, 830845 (US). Bellenden Ker: 1889, Bailey '545' (BRI); Centre peak, 7 km W of 

Bellenden Ker, A. & M. Aptroot 22485, 22499, 22502, 22520 (ABL). Atherton Tablelands: Davies Creek Rd. 17 

km S of Kennedy Hwy., S of Davies Creek Falls NP., E of Mareeba, Hale 832090 (US); Plath Rd. logging head, 

9 km W of Plath Rd., off Kennedy Hwy, Herberton Range, S of Atherton, Hale 831942, 832088, 832100, 

832438 (US); Tumoulin Rd., 5 km from turnoff to Ravenshoe, Lumbsch & Mangold 19151 zb, Mangold 30 zk, 

zl, zp, zq, zu (F). Dawson logging area, State Forest Reserve 605, 24 km S of Koombooloomba turnoff, WSW of 

Tully, Hale 832651, 832653 (US). Culpa logging area, 13 km from Koombooloomba rd. turnoff, SE of Tully 

Falls, Hale 832383, 832552 (US). Tully Gorge NP., Ewers 7887 (CANB). 20 km on the Kirrama Forest Rd., W 

of Kennedy, Hale 830253 (US). Crystal Creek NP., Paluma, Rambold 4779 (M). Mt. Spec NP., Ridge on the 

Loop, on the Paluma Road, WNW of Townsville, Hale 830716, 832368, 832422, 832548, 832763 (US). 

Eungella NP.: Along Broken River, Lumbsch & Mangold 19100 k, y (F); Trail from Broken River station to 

rainforest, Lumbsch & Mangold 19108 h, j (F); Rd. to rainforest at NP. margin, Lumbsch & Mangold 19110 b 

(F). Kondalilla Falls, SW of Nambour, Hale 831987 (US). Maleny Range, Mary Cairn Cross Park about 6 km S 

of Maleny, Hafellner 16676 (GZU), Hale 831460, 831720 (US). Kenilworth SF., SW of Kenilworth, Hale 

831687 (US). Booloumba Creek SF., SW of Kenilworth, Hale 831253 (US). Amama SF., SW of Kenilworth, 

Hale 830303, 831191, 832517 (US). D'Anguilar Range NW of Brisbane, W of Mt. Glorious township, Hafellner 

16974 (GZU). Mt. Mee SF, near Mt.Mee, N of Brisbane, Hafellner 16920 (GZU), Rogers 2418, 2499 (BRI), 

2498 (BRI, GZU), Hale 53553, 53576, 53581, 53600, 53621, 53641, 58664, 58669, 58696, 58672, 58680, 

830837, 83-2370, 831271, 831610, 832366, 832407 (US). C. 10 km NE Killarney, Hale 58836 (US). Carabeen 

Nature Refuge, 45 km E of Warwick, Lumbsch & Mangold 19174 i, x, 19175 i (F). Head of Teviot Brook, NE of 

the Head, Boonah rd., Hale 59481, 59482 (US). 13 km N of O'Reilly's on rd. to Lamington NP., Hale 831848, 

831916, 832077, 832371, 832372, 832420, 832750 (US). Lamington NP.: Mc Pherson Range: Binna Burra, 

Track from Car Park to Tullawallal, Lumbsch & Dickhaeuser 11011 f (F), Duck Creek Rd., K. & A. Kalb 34273 

(hb. Kalb); Python Rock Track, Hale 830941, 831317, 832081, 832699 (US); O'Reillys Guesthouse, K. & A. 

Kalb 21537, 21544 (hb. Kalb); Main Border Track out of O'Reillys, Hale 830963, 832268 (US); Moran Falls, 

Tibell 12682 a (UPS). New South Wales: Mt. Warning NP.: Hale 831869, 832373, 832374, 832439 (US); Track 

from summit to parking lot, Mangold 19 i, m (F). Lions Tourist Rd. nr. Queensland border, N of Waingaree,


Hale 830836, 832461 (US). Border Ranges NP., NE of Wiangaree, Brindle Creek, Hafellner 19364 (GZU). 

Nightcap NP., Mt. Nardi/Mt. Matheson Track, Mangold 22 c, zc (F). Nightcap Forest Drive, W of 

Mullumbimby: Gibbergunyah Roadside Reserve, Whian Whian SF., Hale 830122 (US); Big Shrub Flora 

Reserve, Hale 831378 (US). Big Shrub, Richmond River, Jul. 1894, Wilson s.n. (NSW-153760, -539416). Cattle 

Creek SF., Briggsvale, 12 km NNE of Dorrigo, Verdon 3837 (CANB, H). Dorrigo NP., Never Never Picnic Area 

and Rosewood Creek Track, Mangold 24 l (F). Doyles River SF. on Oxley Hwy. 95 km SE Walcha, Hale 58582, 

58678, 58893 (US). Bulahdelah District, Myall River SF., E von Stroud, Jarrah Rd, Kalb & Filson 17930, 17957 

(hb. Kalb). Along Mill Creek, c. 50 km NW von Sydney, Kalb & Archer 33839 (hb. Kalb). Hornsby, Sept. 1897, 

Wilson s.n. (NSW-539363, -603866). Near Blackheath, Blue Mountains, Hale 58575, 58583, 58599, 58611, 

58659 (US). Below Katoomba Falls-trail to Giant Stairway, Katoomba, Hale 58719 (US). Royal NP., Bola 

Creek, E of Waterfall, K. & A. Kalb 20555, 21584, 21731 (hb. Kalb). Lord Howe Island, Track from Smoking 

Tree Ridge to Rocky Run, Elix 42459 (CANB). Sri Lanka: Central Province, Hale 50268 (US). Sabaragamuwa 

Province, Hale 51208 (US). 

Thelotrema oleosum Mangold spec. nov. ined. 

Type: Australia, New South Wales, Dorrigo NP., Sassafras Creek Track, Mangold 25 m (CANB-holotype, 

NSW-isotype). 

ETYMOLOGY – The epithet refers to the inspersed hymenium of the species (from lat. 

oleosus =oily). 



greenish-gray to (pale) olive, with a distinct reticulate pattern. Surface dull to slightly shiny, 

smooth, usually distinctly verrucose to verruculose, ±fissured. Thallus cover variable, often 

with an incontinuous to continuous true cortex consisting of periclinal hyphae up to c. 30 µm 

thick, sometimes cortex structures absent or covered by an incontinuous protocortex. Algal 

layer moderately well developed, often incontinuous due to crystal inclusions, calcium 

oxalate crystals abundant, small to large, often in clusters or more rarely scattered. Vegetative 

propagules not seen. Ascomata inconspicuous, large, up to 1.5 mm in diam., roundish to 

irregular in fused ascomata, perithecioid, solitary to marginally fused, immersed to emergent, 

then hemispherical to somewhat subglobose. Disc not visible from surface. Pores small, up to 

c. 150 µm, roundish to more rarely roundish-irregular, entire, upper parts of proper exciple 

usually visible from surface, fused to free, whitish to grayish, rarely slightly shrunken, 

incurved. Thalline rim margin roundish to roundish-irregular, small, entire, moderately thin to 

moderately thick, incurved, concolorous with thallus to pale brownish. Proper exciple fused, 

becoming apically detached in older stages, moderately thin, hyaline internally, pale 

yellowish and usually with substrate particles incorporated marginally, apically (dark-)brown 

to more rarely slightly carbonized, non-amyloid. Hymenium up to c. 400 µm high, distinctly 

inspersed, strongly conglutinated, paraphyses ±bent and wavy, interwoven, unbranched, tips 

unthickened to slightly thickened, lateral paraphyses present, conspicuous, up to c. 30 µm 

long, appearing coarse, columellar structures absent. Epihymenium indistinct, hyaline, 

without granules. Asci 1-spored, tholus absent, often with somewhat thickened lateral ascus 

walls in younger stages. Ascospores (very) large, densely eumuriform, cell walls and 

endospore thin, often with a very thin, indistinct halo, becoming grayish to brownish in late 

maturity, dark brown in depauperate ascospores, ±distinctly amyloid, predominantly in 

hyaline stages, oblong to fusiform, with roundish to narrowed-roundish ends, loci small, 

roundish to somewhat angular, predominantly irregular, transverse septae thin, ±distinct, 

regular to slightly irregular, 120-250 x 30-50 µm with multiple loci. Pycnidia not seen.


Fig. 150. Thelotrema oleosum: growth habit (A), ascomata (B), ascoma and thallus section (C), 

ascospore showing amyloid reaction (D), detail of ascospore (E) and younger ascospore (F). A.-F.: 

CANB-holotype. Bar= A: 1.5 mm; B: 1 mm; C: 400 µm; D, F: 30 µm; E: 10 µm. 


ECOLOGY AND DISTRIBUTION – Thelotrema oleosum grows on tree bark in (sub)tropical 

rainforests in altitudes ranging from 50 to 1130 m. It is rare but wide-spread known from 

northern and southern Queensland and the Queensland/ New South Wales border region. 

NOTES – This new species is characterized by a verrucose to verruculose thallus with 

reticulate pattern due to the abundance of calcium oxalate crystals, perithecioid ascomata with 

a fused to apically free proper exciple, inspersed hymenium, distinct, coarse lateral 

paraphyses, monosporic asci with large, eumuriform, brownish, amyloid, thin-walled 

ascospores and the absence of secondary metabolites. A similar species is T. saxicola, see


under that species for differences. Thelotrema 

porinaceum is also similar, but can be readily 

distinguished by the presence of norstictic acid and 

the non-amyloid ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Atherton 

Tablelands, Tumoulin Rd., 5 km from turnoff to Ravenshoe, 

Lumbsch & Mangold 19151 x (F). Culpa logging area, 13 km 

from Koombooloomba rd. turnoff, SE of Tully Falls, Hale 

830718 (US). Mt. Spec NP., Ridge on the Loop, on the 

Paluma Rd., WNW of Townsville, Hale 831919, 832396 

(US). Wooroi State Forest Park, W of Teewantin, Hale 

831957 (US). 13 km N of O'Reilly's on rd. to Lamington NP., 

S of Brisbane, Hale 830656, 831928 (US). Lamington NP., 

Main Border Track out of O'Reillys, Hale 831509 (US). New 

South Wales, Dorrigo NP., Never Never Picnic Area and 

Rosewood Creek Track, Mangold 24 a (F). 

Thelotrema pachysporum Nyl. 

Bull. Soc. Linn. Normandie 2(2): 72 (1868). Phaeotrema pachysporum (Nyl.) Zahlbr., Cat. Lich. Univ. II: 

609 (1923). Type: New Caledonia, Pancher s.n. (H-Nyl.22747-holotype). 

Thelotrema exalbidum Stirt., Proc. Phil. Soc. Glasgow 13: 184 (1881). Leptotrema exalbidum (Stirt.) Zahlbr., 

Cat. Lich. Univ. 2: 634 (1923). Type: India, Assam, 03. Feb. 1879, Watt s.n. (BM!-lectotype, selected by Hale 

[1974: 29]; GLAM!-isolectotype). 

Thelotrema galactinum Vain., Dansk Bot. Ark. 4(11): 24 (1926). Phaeotrema galactinum (Vain.) Zahlbr., 

Cat. Lich. Univ. 8: 245 (1932). Type: Mexico, "Malvapam" (=Maloapam), Liebmann 7712 (C!-lectotype, 

selected by Hale [1974: 29]). 

Thelotrema limae Vain., Boletim Soc. Broteriana, sér. 2, 6: 150 (1929). Phaeotrema limae (Vain.) Zahlbr., 

Cat. Lich. Univ. X: 216 (1939). Type: Mozambique, Palma, 1916, A. Pires de Lima 38 (TUR-Vain. 34808holotype). 

Thelotrema palmense Vain., Boletim Soc. Broteriana, sér. 2, 6: 150 (1929). Phaeotrema palmense (Vain.) 

Zahlbr., Cat. Lich. Univ. X: 216 (1939). Type: Mozambique, Palma, 1916, A. Pires de Lima 78 (TUR-Vain. 

34810-lectotype, selected by Salisbury [1972: 270]). 



T. oleosum.. 

Thallus predominantly to entirely hypophloedal, up to c. 80 µm high, pale gray to pale 

yellowish-gray. Surface dull to rarely slightly shiny, rough, continuous, unfissured to 

sometimes slightly fissured. Cortex structures absent. Algal layer incontinuous and poorly 

developed, calcium oxalate crystals abundant, small. Vegetative propagules not seen. 

Ascomata inconspicuous, (moderately) large, up to c. 1 mm in diam., apothecioid, roundish, 

solitary, immersed to usually ±emergent, predominantly hemispherical. Disc usually partly 

visible from surface, grayish, coarsely pruinose. Pores moderately wide to gaping, up to c. 

700 µm in diam., roundish to slightly irregular, apical to upper proper exciple visible from 

surface, free, predominantly entire to rarely slightly jagged, whitish to off-white, 

predominantly incurved, sometimes slightly shrunken. Thalline rim margin moderately thin, 

roundish, entire to slightly split, in older ascomata often becoming eroded, concolorous with 

thallus to brownish in eroded ascomata due to protruding substrate layer, thalline rim 

concolorous to grayish brown in specimen with thinner thallus, with same surface as thallus, 

incurved. Proper exciple free, moderately thin, hyaline internally, yellowish-brown to


grayish-brown or dark-brown marginally, sometimes with incorporated small calcium oxalate 

crystals, rarely faintly amyloid at the base. Hymenium up to c. 200 µm high, non-inspersed, 

moderately conglutinated, paraphyses slightly bent, slightly interwoven, unbranched, tips 

slightly thickened, somewhat irregular, lateral paraphyses present, sometimes inconspicuous, 

up to c. 25 µm long, columellar structures absent. Epihymenium moderately thick, hyaline, 

with grayish to brownish granules, sometimes small crystals and remnants of deceased 

ascospores. Asci 4-8-spored, tholus thick, moderately thick at maturity. Ascospores 

moderately small to moderately large, transversely septate, very rarely with a single, thin 

longitudinal septum, cell walls thick to very thick, halonate, brown, moderately to weakly 

amyloid, clavi- to fusiform, with narrowed-rounded to subacute ends, loci roundish, 

predominantly oblong to lentiform, septae moderately thin to moderately thick, regular, 30-75 

x 10-15 µm with 8-18(20)x(2) loci. Pycnidia not seen. 

CHEMISTRY – Thallus K-, C-, PD-; no secondary compounds detectable by TLC (results 

often falsified due to contamination by host substrates). 

Fig. 152. Thelotrema pachysporum: growth habit (A), ascomata (B), mature ascospore (C) and 

younger ascosporez (D). A.-D.: Stevens 4109. Bar= A: 1.5 mm; B: 1 mm; C: 10 µm; D: 12 µm.



pachysporum grows on bark in mangroves and 

sclerophyll forests, grasslands or rarely rainforests. 

The species was collected in altitudes ranging from 

sea level up to 780 m. It is a moderately common 

and wide-spread species in the north-western 

Northern Territory and Queensland. This is the first 

report for Australia. It has previouly been recorded 

from the Neotropics (Frisch, 2006), New Caledonia, 

India and Africa, indicating a pan(sub)tropical 

distribution. 


ecorticate thallus, large ascomata with free proper 

exciple, moderately large, transversely septate, 

thick-walled, brown ascospores with roundish loci 

and the absence of secondary compounds. Besides 

T. lacteum (see below) the only similar species is T. 

rockii from Hawaii, which is distinguished by a 

higher hymenium (up to 250 µm), larger ascospores 

(up to 85 µm long) with a weak amyloid reaction (non-amyloid to faintly amyloid in 

immature stages) and the occurrence of the stictic acid chemosyndrome. Unfortunately the 

type of T. pachysporum was not available for study, however, Frisch (2006) provided a 

detailed description. Authors published different interpretations of the taxonomy of this 

species. Frisch (2006) considered T. pachysporum as a distinct species from the similar T. 

lacteum (for differences see under that species) where it was recently synonymized by Hale 

(1981). Salisbury (1972) erroneously reported transversely septate ascospores in the FH- 

Tuck. type of T. lepa-dodes (see also Hale, 1978), and consequently placed several names 

synonymous which belong to T. pachysporum (T. exalbidum, T. limae and T. pal-mense). 

Hale (1981) referred to additional syno-nyms for T. lacteum in his publication (Hale, 1974) 

for Phaeotrema disciforme (=T. monosporum), where three names are listed: T. exalbidum 

and T. galactinum (= T. pachysporum syn. nov.), and T. aquilinum (= T. monosporum). 

SPECIMENS EXAMINED – Australia, Northern Territory: 7.5 km E of Darwin, 2.5 km N of Landing Quarantine 

station, Thor 5926 (S). 42.5 km ESE of Darwin, Black Jungle, Thor 5779 (S). Queensland: At Musgrave, on 

Peninsula Developmental Rd., Cape York Peninsula, Hale 831172 (US). Cape Tribulation Area, Myall Beach, 

Lumbsch & Mangold 19158 l, 19162 d, e, j, k (F). Cooktown Rd., 3 km NW of Mt. Molloy, Elix & Streimann 

17163 (CANB). Ravenshoe SF., Culpha Ck. Catchment, Cardwell Range, 41 km SE of Ravenshoe, Elix & 

Streimann 16091 (CANB). Butchers Hill Property, 6 km from homestead, Stevens 4109 (BRI). Babinda 

Boulders, Lumbsch & Guderley 11151 m (F). Cape Cleveland NP., E of Townsville, Hale 831747 (US). First 

Turkey, Mt. Archer Environmental Park, 7 km NE of Rockhampton, Elix 34535 (B, CANB). Tandora, about 25 

km ENE of Maryborough, Hafellner 18267, 18269 (GZU). Imbil SF., 6 km NW of Imbil, Rogers 2536 (BRI). 

Thelotrema polythecium Nagark., Sethy & Patw. 

Mycotaxon 28: 195 (1987). Type: India, Andaman Islands, North Andaman, Mayabandar Range, Nagarkar 

& Patwardhan 852793 (AMH-holotype, US-isotype - not seen). 



T. pachysporum.


Fig. 154. Thelotrema polythecium: ascomata (A, B) and ascospores (C-E). A.-E.: Hale 69145. Bar= 

A: 0.25 mm; B: 0.2 mm; C: 3 µm; D: 4 µm; E: 6 µm. 

Thallus hypophloedal, pale greenish-gray. Surface dull to slightly glittering, porous due to 

visible substrate structure, continuous, unfissured. Cortex structures absent. Algal layer 

poorly developed, incontinuous, calcium oxalate crystals not seen. Vegetative propagules not 

seen. Ascomata inconspicuous, moderately large, up to c. 500 µm in diam., roundish to 

somewhat irregular in fused ascomata, becoming apothecioid to somewhat chroodiscoid in 

older stages, erumpent, solitary to marginally fused, sometimes clustered in small groups, 

immersed. Disc becoming partly to rarely entirely visible from surface, pale yellowish to pale 

flesh-colored, epruinose. Pores becoming wide to gaping, up to c. 400 µm in diam., roundish 

to irregular, split, proper exciple not visible from surface. Thalline rim margin (moderately) 

thick, lacerate to coarsely squamose, sometimes slightly layered, whitish to off-white, 

incurved to somewhat erect in outer parts. Proper exciple fused, moderately thin to 

moderately thick, often indistinct due to incorporated substrate layers, hyaline internally, pale 

yellowish-gray marginally, non-amyloid. Hymenium up to c. 90 µm high, non-inspersed, 

weakly conglutinated, paraphyses parallel, predominantly straight, unbranched, tips 

unthickened to slightly thickened, lateral paraphyses present, inconspicuous, up to c. 20 µm 

long, often not clearly separated from proper exciple, columellar structures absent. 

Epihymenium indistinct, hyaline to slightly grayish, without granules or with fine hyaline to 

pale grayish granules. Asci 8-spored, tholus thin, not visible at maturity. Ascospores small, 

(sub-)muriform, cell walls moderately thin, endospore moderately thick, with thin halo, 

hyaline, non-amyloid, young ascospores claviform, becoming oblong to roundish-claviform at 

maturity with roundish to narrowed-roundish ends, loci small, distinctly angular and irregular 

in younger stages, roundish to slightly angular and subglobose to oblong to somewhat 

irregular at maturity, with same shaped to hemispherical (or in young stages conical) end


cells, transverse septae moderately thick, distinct, regular, 17-25 x 6-9 µm with 6-9 x 1-3(4) 




polythecium is known in Australia from one 

collection on tree bark in a tropical rainforest at an 

unknown altitude in northern Queensland. This is 

the first report for Australia of a species that was 

previously only known from the Andaman Islands. 

NOTES – Although the type material of this taxon 

was not available for study, the singular Australian 

specimen agrees well with to the original 

description of T. polythecium (Nagarkar & al., 

1987). It has an hypophloedal, ecorticate thallus, 

somewhat chroodiscoid ascomata with a bright, 

lacerate to coarsely squamulose margin, typical, 

small, (sub-)muriform, hyaline, non-amyloid 

ascospores with distinctly thickened endospore and 

lacks secondary compounds. Thelotrema subadjectum 

and T. cyphelloides are similar; see there 

for differences. 

The generic position of T. polythecium is 

uncertain. The ascospores are similar to the O. clandestina-group, however, the lack of 

carbonization and the presence on lateral paraphyses differentiate it from that species group. 

The hymenium and with some respects the excipular structures as well as the thalline rim 

morphology are similar to Topeliopsis, the immersed, ±chroodiscoid ascomata and the lack of 

an amyloid reaction of the ascospores, on the other hand resemble Chapsa. Molecular data 

will be necessary to re-evaluate the generic placement of this species. 

SPECIMENS EXAMINED – Australia, Queensland: Wooroonooran NP. ("Bartle Frere NP."), Trail head, Hale 

69145 (US). 

Thelotrema porinaceum Müll.Arg. 

Nuov. Giorn. Botan. Ital. 23: 130 (1891). Myriotrema porinaceum (Müll.Arg.) Hale, Mycotaxon 11: 134 

(1980). Type: Japan, Awa, Miyoshi 17 (G!-holotype). 



T. polythecium. 

Thallus corticolous or rarely saxicolous, predominantly episubstratic, (moderately) thick, 

up to c. 800 µm high, pale grayish-green to (pale) olive. Surface dull to shiny, smooth to 

rarely somewhat roughened, continuous to usually verruculose or verrucose, ±distinctly 

fissured. Cortex structures absent or covered by an incontinuous protocortex up to c. 20 µm 

thick. Algal layer variable, mostly continuous and well developed, sometimes discontinuous 

and poorly developed, calcium oxalate crystals abundant, large, scattered to clustered. 

Vegetative propagules not seen. Ascomata predominantly inconspicuous, (moderately) large, 

up to 1.2 mm in diam., roundish, solitary, perithecioid, immersed to more often ±distinctly 

emergent, then predominantly (verrucose-)hemispherical to rarely somewhat subglobose.


Fig. 156. Thelotrema porinaceum: growth habit (A), ascomata (B, C) and ascospores (D, E). A.: 

Mangold 19 g; B.: G-holotype; C.-E.: Lumbsch & Mangold 19108 d. Bar= A: 10 mm; B: 1 mm; C: 

0.7 µm; D: 20 µm; E: 25 µm. 

Disc not visible from surface. Pores tiny to small, up to c. 80 µm in diam., roundish, entire, 

apical proper exciple becoming visible from surface, forming a fused to entirely free inner 

pore margin, pore margin level with thallus to more often sunken, off-white to pale brownish, 

incurved. Thalline rim margin moderately thick, roundish, entire, concolorous with thallus or 

somewhat brighter, level with thalline rim or ±sunken, thalline rim not developed or not 

distinguishable from thallus in immersed ascomata, in emergent ascomata incurved, with 

same surface as thallus. Proper exciple fused to apically or in upper parts free, hyaline to pale 

yellowish internally, yellowish-gray to yellowish-brown marginally, ±distinctly amyloid at 

the base. Hymenium up to c. 400 µm high, either non-inspersed (strain I) or densely inspersed 

(strain II), moderately conglutinated, paraphyses thin, distinctly bent to curly, particularly 

towards the tips, parallel to slightly interwoven, unbranched, tips unthickened, lateral 

paraphyses present, usually inconspicuous,not conglutinated, up to c. 35 µm long, columellar 

structures absent. Epihymenium indistinct, hyaline, without granules or crystals. Asci 1- to 

very rarely 2-spored, tholus indistinct, lateral walls thickened, thin when mature. Ascospores 

(very) large, densely eumuriform, cell walls and endospore thin, non-halonate, hyaline, rarely 

pale yellowish in older stages, non-amyloid to faintly amyloid in mature spores, 

predominantly oblong-fusiform, often irregular in outline or bifusiform, straight to slightly 

bent, ends roundish to predominantly narrowed-roundish, loci angular to slightly roundish,


predominantly irregular, transverse septae thin, becoming somewhat indistinct with age, 

regular to slightly irregular, 100-230 x 20-45 µm with multiple loci. Pycnidia not seen. 




porinaceum occurs on bark or rarely siliceous rocks 

in tropical to warm-temperate rainforests and wet 

sclerophyll forests in altitudes ranging from 100- 

1200 m. This common and widespread species 

occurs along the east coast east of the Great 

Dividing Range from northern Queensland to 

southern New South Wales. It is also known from 

Sri Lanka (Hale, 1981) and Japan, being a 

paleosubtropical element. 

NOTES – This taxon is readily distinguished by a 

thick, ecorticate thallus, perithecia, large eumuriform, 

predominantly colorless ascospores and 

the norstictic acid chemosyndrome. A large group 

of otherwise identical specimens (over 100 

collections in Australia, here referred to as strain II) 

differ from the type material and the remaining 

Australian samples (strain I) in a strongly inspersed 

hymenium. Since the nature of hymenial inspersion 


T. porinaceum. 

as a species delimiting character is not well understood this form is treated here as a strain 

within the species. Additional molecular data will be necessary to evaulate the taxonomic 

importance of this character. Similar, norstictic acid containing species include: T. 

eungellaense (for differences see under this species), Myriotrema frustillatum, T. weberi and 

T. subweberi. The latter two taxa, can be distinguished by a thinner thallus, distinctly 

apothecioid ascomata and thick-walled, distinctly amyloid ascospores. Myriotrema 

frustillatum lacks lateral paraphyses and is isidiate. Other similar species (particularly to the 

strain II specimen), with a different chemistry are T. oleosum and T. saxicola, see under these 

species for differences. 

SPECIMENS EXAMINED – Strain I: Australia, Queensland: Cedar Bay NP., Track to Mt. Finnigan, Streimann 

57062A (CANB). Thornton Range, NW of Mossman, Hale 830965, 831306, 831405, 832135, 832362 (US). Mt. 

Windsor, NW of Mossman, Hale 830620, 830658, 832448, 832560, 832710, 832711 (US). Daintree NP., 

Mossman sct., lower Mossman River, Mangold 36 a, l (F). Mt. Lewis Rd., W of Mossman, Hale 831238, 

831843, 831945 (US). Kuranda, 1893, Wilson s.n. (H). Davis Creek Rd., E of Mareeba, Hale 830668 (US). 

Atherton Tablelands: Lake Tinaroo, Lumbsch & Mangold 19125 d, p (F); Tumoulin Rd., 5 km from turnoff to 

Ravenshoe, Lumbsch & Mangold 19148 m, 19151 w (F); Souita Falls, Lumbsch & Mangold 19155 t, 19156 d, g, 

h (F); Plath Rd. logging head, Herberton Range, Hale 64187, 830645, 830657, 830660, 830675, 831844 (US); 

Danbulla Forest Drive, Hale 832304 (US); Mt. Hypipamee NP., Hale 832678 (US); 13 km S of Ravenshoe on 

Tully Falls Rd., Hale 830730, 832667, 832668 (US). Babinda Boulders, Mangold 39 q, ze, zf (F). Francis Range, 

NW of Innisfail, Hale 832217 (US). Mt. Chalmynia logging area, W of Innisfail, Hale 832347 (US). Culpa 

logging area, SE of Tully Falls, Hale 830652, 832690 (US). Mt. Tyson track, 2 km W of Tully, Streimann 45617 

(CANB). Kirrima SF., Cardwell Range, Elix 15686 (CANB). 8 km E of Wallaman Falls, W of Ingham, Hale 

830702, 832358 (US). Little Crystal Creek Falls-Mt. Spec NP., Hale 831471 (US). Murray Falls, W of Kennedy, 

Hale 831306, 831340, 831426, 831449, 831529 (US). Eungella NP.: Trail from Broken River station to 

rainforest, Lumbsch & Mangold 19108 d, j (F); Near Peases Lookout, Hale 831422, 831755 (F). Softwood shrub 

near Yabba Rd., 6 km N of Jimna, Rogers 2269 (BRI). Lamington NP.: O'Reillys Guesthouse, K. & A. Kalb 

21541 (hb. Kalb); Phyton Rock Track, Hale 830630 (US). Carabeen Nature Refuge, 45 km E of Warwick,


Lumbsch & Mangold 19175 c (F). New South Wales: Lions Tourist Rd., N of Waingaree, Hale 830642 (US). 

NW part of Wiangaree Forest Drive, N of Kyogle, Hale 831232, 831301, 831318, 832302 (US). Mt. Warning 

NP., Mangold 19 p (F). Dorrigo NP., Mangold 26 a (F). Dorrigo SF., Hale 58738, 58750, 58770 (US). Doyles 

River SF., 95 km SE of Walcha, Hale 58542, 58578, 58710 (US). Barrington Tops NP., NE von Scone, K. & A. 

Kalb 18025 (hb. Kalb). Bulahdelah District, Myall River SF., K. & A. Kalb 18051, 18063, 18065 (hb. Kalb). 

Trail along bank of Mill Creek, 50 km NW of Sydney, K. & A. Kalb 27048 (hb. Kalb). Track by side of Magdala 

Creek, 1 km S of Springwood, Archer P20 (HOB). Glenbrook Creek, SSE of Springwood, K. & A. Kalb 21885, 

21886 (hb. Kalb). Tianjara Falls, 30 km NW of Ulladulla, Streimann 7857 (H, CANB). 

Strain II: Australia, Queensland: Mt. Windsor logging area, NW of Mossman: near jct. rd. to old Forestry 

Camp and main rd., Hale 830626, 830669 (US); E of old Forestry Camp, Hale 830949, 832715 (US). Mt. Lewis 

Rd., 4 km N from Kennedy Hwy., W of Mossman, Hale 832097 (US). End of Clohesy River Rd. c.16 km SE 

Kennedy Hwy., W of Cairns, Hale 830638 (US). Atherton Tablelands: Lake Tinaroo, Downfall Creek Camping 

Area, Lumbsch & Mangold 19123 m (F); Lake Barrine NP., Hale 831597 (US). Lake Eacham NP., Mangold 29 

t, bj (F); Plath Rd. logging head, 9 km W on Plath Rd., off Kennedy Hwy, Herberton range, S of Atherton, Hale 

832157 (US); 18 km S of Atherton, Mt. Hypipamee, at Crater Lake, Tibell 15431 (UPS); Area below crater, Mt. 

Hypipamee NP., Hale 831913, 832221 (US); Tumoulin Rd., 5 km from turnoff to Ravenshoe, Lumbsch & 

Mangold 19151 zb, Mangold 30 p (F); Souita Falls, Lumbsch & Mangold 19155 r (F). State Forest area on Tully 

Rd., 1 km from jct. with S. Mission Beach Rd., S of Mission Beach, Hale 831338, 831508, 831510, 831538, 

831730, 831735 (US). 7.5 km E of Wallaman Falls, W of Ingham, Hale 831941 (US). Dawson logging area, 24 

km S of Koombooloomba turnoff, WSW of Tully, Hale 830690, 830694, 832701 (US). Little Crystal Creek 

Falls-Mt. Spec NP., the falls on Paluma Rd., Hale 831743 (US). Eungella NP.: Rosser Rd. entry point off 

Darymple rd., near Peases Lookout, Hale 831250, 831436 (US). Waterpark Creek State Forest Park in Byfield 

SF., 24 km N of Yeppoon, Hale 831511 (US). Amama State Forest Park, 6 km SW of Amamoor, S of Glympie, 

Hale 832508 (US). Booloumba Creek SF., SW of Kenilworth, Hale 831214 (US). Mt. Mee SF., nr. Mt. Mee, N 

of Brisbane, Hale 58562, 58589, 58612, 58681, 58689, 832332 (US). Mt. Tennison Woods, near Mt. Glorious, 

Rogers 694624/5 (BRI). Mt. Glorious: Brisbane SF., K. & A. Kalb 34281 (hb. Kalb); Bennet Rd., Rogers 8447, 

8448 (BRI). D'Anguilar Range NW of Brisbane, W of Mt. Glorious township, Hafellner 16980, 16986 (GZU). 

Upper Coomera, Wilson s.n. (NSW-539334, -539336). Cameron falls NP., Elix 1127 (CANB). Cunninghams 

Gap NP., 50 km NE of Warwick, Hale 66699, 831749, 831757, 831758 (US). Lions Tourist Rd. near 

Queensland border, N of Waingaree, Hale 69161 (US). Head of Teviot Brook, NE of the Head, Boonah road, 

Hale 59483 (US). Lamington NP.: Phyton Rock Track, Hale 830653, 830979, 830984, 832642 (US); O'Reillys 

Guesthouse, K. & A. Kalb 21526 (hb. Kalb); Moran Falls, Tibell 12684 (UPS); Main Border Track out of 

O'Reillys, Hale 832270 (US); Along Blue Pool Track, Tibell 12668 (UPS); Along circular track, Foresters 

Camp, Lyrebird Lookout, Moonlight Crag, Brownlie 1023644 pr. p. (MEL). New South Wales: Tooloom NP., 

23 km WSW of Woodenbong, Streimann 60911A (CANB). Lions Tourist Rd. near Queensland border, N of 

Waingaree, Hale 830635, 830636, 830672, 830829, 830861, 830880, 831940, 831948, 831959, 832709, 832735, 

832741, 832777 (US). Victoria Park Nature Reserve, SE of Lismore, S of Bruxner Hwy., Hale 831333 (US). 

McPherson Range, Border Ranges NP., NE of Wiangaree, uppermost part of Gradys Creek, Hafellner 16661 

(GZU). NW part of Wiangaree Forest Drive, N of Kyogle, Hale 831309, 831331, 831332 (US). Mt. Warning 

NP., W of Murwillumbah: Hale 83041, 83054, 831613, 832069, 832694 (US); track from summit to parking lot, 

Mangold 19 g, k, n, zc (F). Tweed Range, Mebbin NP., 25 km SW of Murwillumbah, Mangold 21 a (F). 

Nightcap Forest Drive, W of Mullumbimby: Whian Whian SF., Hale 830116, 830120, 830123, 831219, 831737 

(US); Big shrub Flora Reserve, Hale 831299, 831404 (US). Nightcap NP., Mnt. Nardi/Mnt. Matheson Track, 

Mangold 22 e, v, z, za (F). Cambridge Plateau Forest Drive, N of picnic area, Richmond Range SF., 30 km W of 

Casino, Hale 830117, 830118, 831149, 831158, 831304, 831682, 832546 (US). Wollongbar, Jun. 1894, Wilson 

s.n. (NSW-539335, -539382). Big Shrub, Richmond River, Jul. 1894, Wilson s.n. (NSW-539337). Cattle Creek 

SF., Briggsvale, 12 km NNE of Dorrigo, Verdon 3832 (B, US). Dorrigo NP., Sassafras Creek Track, Mangold 

25 j (F).


Thelotrema porinoides Mont. & Bosch 

in Miguel, F. A. W., Plant. Jungh. fasc. 4: 484 (1855). Ocellularia porinoides (Mont. & Bosch) Zahlbr., Cat. 

Lich. Univ. II: 599 (1923). Thelotrema cavatum var. porinoides (Mont.) Nyl., Mém. Soc. Bot. Cherbourg 5: 118 

(1858). Type: Java, Junghuhn s.n. (L-lectotype, selected by Hale [1974b: 20]; FH-Tuck-, G!-, H-Nyl. 22732-, 

PC-, W-isolectotypes). 

Thelotrema albidiforme Leight., Trans. Linn. Soc. London 27: 170 (1869). Ocellularia albidiforme (Leight.) 

Zahlbr., Ann. Mycol. 14: 50 (1916). Type: Sri Lanka, Thwaites 19 (BM!-holotype; H-Nyl.22733-, PDAisotypes). 

Thelotrema exanthismocarpum Leight., Trans. Linn. Soc. London 27: 169 (1869). Ocellularia 

exanthismocarpa (Leight.) Zahlbr., Cat. Lich. Univ. II(4): 590 (1923). Type: Sri Lanka, Thwaites 97 (BM!lectotype, 

selected by Hale [1974: 21]). 

Thelotrema obovatum Stirt., Trans. Glasgow Soc. of Field Natural. 1: 21 (1873). Ocellularia obovata (Stirt.) 

Müll. Arg., Bull. Herb. Boissier 2, append. 1: 74 (1894). Type: New Zealand, Wellington, Buchanan 57b 

(GLAM!-lectotype, selected by Galloway [1985: 576]; BM!-isolectotype). 

Ocellularia platychlamys Müll. Arg., Bull. Herb. Boissier 3: 313 (1895). Type: Australia, Queensland, 

Blackall Range, Eumundi ['Emundi'], Bailey s.n. (G!-lectotype, here selected). 

Thelotrema homothecium Vain., Ann. Acad. Sci. Fenn. A 15(6): 190 (1921). Ocellularia homothecia (Vain.) 

Zahlbr., Catal. Lich. Univ. 2: 593 (1923). Type: Philippines, Irosin, Elmer 14852 pr.p. (TUR-Vain.-holotype, 

US!-isotype). 

[For additional synonyms see Frisch, 2006.] 


Thallus predominantly hypophloedal, very thin, epiphloedal parts (usually in ascomata area) 

up to c. 30 µm high, in shades of grayish-green. Surface dull to slightly shiny, smooth to 

roughened due to protuberant substrate, continuous to slightly verruculose, unfissured 

(sometimes appearing fissured due to substrate structure), covered by continuous to 

incontinuous protocortex, up to c. 30 µm thick. Algal layer variable, well developed, 

continuous to incontinuous, occurrence of calcium oxalate crystals variable, usually frequent, 

small to large, often clustered. Vegetative propagules not seen. Ascomata ±conspicuous, 

(moderately) large, up to c. 900 µm in diam., roundish, apothecioid, sessile, solitary to 

sometimes fused, usually distinctly emergent, rarely subglobose with constricted base to more 

often hemispherical to broad-cylindrical or cone-shaped with flattened apex. Disc not visible 

from surface to sometimes becoming partly visible, pale grayish, distinctly pruinose. Pores 

small to moderately wide, up to c. 300 µm in diam., roundish, entire to split, proper exciple 

entirely to apically visible from surface, free, apically bright, darker towards the base, 

incurved to slightly erect, sometimes slightly shrunken. Thalline rim margin thick, ±roundish, 

entire to split to lacerate or eroded, often slightly to distinctly layered, incurved to erect, 

mostly whitish or brighter than thallus, sometimes slightly pruinose, thalline rim surface often 

irregular-verrucose to verruculose. Proper exciple free, ±thin, hyaline internally, grayishbrown 

marginally, often covered by grayish granules, non-amyloid to faintly amyloid at the 


interwoven, unbranched, tips slightly thickened, lateral paraphyses present, conspicuous, up 

to c. 40 µm long, columellar structures absent. Epihymenium thick, hyaline, with grayish to 

brownish granules and small crystals. Asci 8-spored, tholus (moderately) thick, not visible at 

maturity. Ascospores typical, variable in size, moderately small to large, transversely septate, 

cell walls very thick, thinly halonate, hyaline, strongly amyloid, oblong-fusiform to 

sometimes oblong-claviform, ends narrowed-roundish to more rarely subacute, loci roundish 

to slightly angular in younger stages, subglobose to predominantly lentiform, end cells 

±conical, septae moderately thin to thick, regular, 40-150(170) x 15-22 µm with 14-26(28) 

loci. Pycnidia not seen, reported by Frisch 2006, see there for description.



stictic (majors), hypostictic (minor), hypoconstictic, cryptostictic and α-acetylhypoconstictic 


Fig. 158. Thelotrema porinoides: growth habit (A, B), ascomata (C, D), ascoma section (E), 

ascospore (F) and ascospore showing amyloid reation (G). A.: GLAM-lectotype of T. obovatum; 

B.: US-isotype of T. homothecium; C., F., G.: Lumbsch & Mangold 19153 w; D.: Lumbsch & 

Mangold 19113 s; E.: Lumbsch & Mangold 19156 b. Bar= A: 1 mm; B: 0.85 mm; C: 0.4 mm; D: 

0.35 mm; E: 100 µm; F: 13 µm; G: 11 µm.



porinoides grows on tree bark in (sub)tropical 


This moderately common and wide-spread species, 

occurs along the east coast of Queensland. The 

pantropical species is also recorded from Hawaii 

(as T. multilocularis – Zahlbruckner, 1912), the 

Neotropics (Hale, 1978b), Africa (Frisch, 2006), 

India (Hale, 1981), Sri Lanka, Japan (Matsumoto, 

2000), Philippines, Borneo (Sipman, 1993), Java, 

Solomon Islands (Hale, 1981) and New Zealand. 


hypophloedal thallus, ±conspicuous, emergent 

ascomata with free proper exciple and a bright, 

usually split to lacerate and often layered thalline 

rim margin, transversely septate, hyaline, thickwalled, 

strongly amyloid ascospores and the 

presence of the stictic acid chemosydrome. Among 

Thelotrema species with transversely septate asco- 

spores and stictic acid, it is characterized by large ascospores. Thelotrema agasthiensis from 

India (type not seen, fide Nagarkar & al. 1988 with ascospores up to 120 µm long with up to 

30 loci), is similar and probably conspecific with T. porinoides. Thelotrema capetribulense 

(ascospores up to 50 µm with up to 14 loci) differs in having a thicker, distinctly verrucose 

thallus and less intensely amyloid, claviform ascospores. Further morphologically similar, but 

chemically differing species include T. nureliyum (secondary compounds absent) and T. 

nostalgicum (protocetraric acid chemosyndrome). Both species are further distinguished by 

larger ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Big Tableland, 26 km S of Cooktown, Elix 17251 (CANB). 

Near Cedar Bay NP., on rd. to Cooktown, Mangold 34 b (F). Cape Tribulation area, near Cape Tribulation Store, 

Mangold 33 o (F). Daintree NP., Mossman Gorge Section, near Rex Creek Swing Bridge, Mangold 35 r (F). End 

of Clohesy River Rd. 16 km SE Kennedy Hwy., W of Cairns, Hale 832265 (US). Crystal Cascades, 5 km W of 

Cairns, Lumbsch & Mangold 19118 e (F). Goldsborough Valley State Forest Park, SW of Gordonvale, Hale 

831973 (US). Atherton Tablelands: Millaa Millaa falls, Lumbsch & Mangold 19137 i (F); 2 km N of S. 

Johnstone Forestry Camp, SE of Millaa Millaa, Hale 832252 (US); Souita Falls, Lumbsch & Mangold 19153 k, 

w, x, 19155 b, w, 19156 b (F). Babinda Boulders, NW of Babinda: Hale 831487 (US); Mangold 39 zg (F). Mt. 

Spec NP., Ridge on the Loop, on the Paluma Rd., WNW of Townsville, Hale 830936 (US). Eungella NP., Finch 

Hatton Gorge, Lumbsch & Mangold 19113 s (F). 

Thelotrema pseudosubtile Mangold spec. nov. ined. 

Type: Australia, Queensland, Crystal Cascades, 5 km W of Cairns, Lumbsch & Mangold 19117 k (CANBholotype, 

BRI-isotype). 

ETYMOLOGY – The epithet refers to the similarities of the new species to T. subtile. 



T. porinoides.


Fig. 160. Thelotrema pseudosubtile: growth habit (A, B), ascomata (C), ascospores (D, E) and 

ascospores showing amyloid reaction (F, G). A.-G.: CANB-holotype. Bar= A: 1 mm; B: 0.5 mm; 

C: 0.3 mm; D, E: 7 µm; F: 12 µm; G: 15 µm. 


gray to pale olive. Thallus dull to slightly shiny, smooth, continuous to ±distinctly verrucose 

to verruculose, slightly to distinctly fissured. Thallus cover variable, thallus covered by an 

incontinuous to continuous protocortex, up to c. 20 µm thick, sometimes becoming distinctly 

conglutinated forming a true cortex of periclinal hyphae up to 40 µm thick. Algal layer 

moderately well developed, continuous, calcium oxalate abundant to sometimes sparse, 

predominantly small to more rarely large, often clustered. Vegetative propagules not seen. 

Ascomata inconspicuous, predominantly small to rarely moderately large, up to c. 300(600) 

µm in diam., roundish, apothecioid, sessile, solitary to marginally or entirely fused, 

predominantly ±immersed to slightly emergent, flattened-hemispherical, with same surface as 

thallus. Disc often becoming partly visible from surface, grayish to pale flesh-colored, 

epruinose to slightly pruinose. Pores (moderately) small, up to c. 150(250) µm in diam., 

roundish to slightly irregular, predominantly entire to slightly split, proper exciple always 

apically to very rarely entirely visible from surface, sometimes somewhat shrunken, whitish 

to off-white, pale brownish towards the base, predominantly incurved. Thalline rim margin


(moderately) thin, becoming moderately wide to rarely gaping with age, roundish to irregularroundish, 

predominantly entire to rarely slightly split, predominantly incurved to more rarely 

somewhat erect, concolorous with thallus or indistinctly brighter. Proper exciple becoming 

partly free to free, predominantly thin, hyaline to pale yellowish internally, brownish or 

grayish-brown marginally, apically often dark-brown, sometimes with substrate inclusions, 

often amyloid at the base. Hymenium up to c. 150 µm high, non-inspersed, distinctly 

conglutinated, paraphyses moderately interwoven, unbranched to sometimes slightly 

branched, tips moderately thickened, lateral paraphyses present, predominantly 


thick, hyaline to sometimes brownish with age, with grayish granules and small crystals. Asci 

2-8-spored, tholus (moderately) thick, thin when mature. Ascospores small to moderately 

large, transversely septate, very rarely with a single longitudinal septae, cell walls 

(moderately) thick, often with distinctly crenate surface, thinly halonate, hyaline, faintly to 

distinctly amyloid, sometimes oblong to fusiform or predominantly clavate, ends roundish to 

acute, loci roundish to somewhat angular, rarely subglobose to more often lentiform or 

slightly irregular, end cells hemispherical to conical, septae moderately thick to moderately 

thin, regular to irregular, 25-60 x 6-9 µm with 7-16 (x 2) loci. Pycnidia not seen. 



pseudosubtile grows on tree bark predominantly in 

(sub)tropical to warm-temperate rainforests, rarely 

in wet sclerophyll forests in altitudes ranging from 

sea level to 950 m. It is common and wide-spread 

occurring in Queensland and New South Wales 

(incl. Lord Howe Island). It is currently only 

known from Australia. 

NOTES – This new species is characterized by a 

thin thallus, predominantly immersed or indistinctly 

emergent, small apothecia with free 

proper exciple, medium-sized, transversely septate, 

weak to moderately amyloid ascospores that are 

hyaline throughout their entire development, have a 

crenate surface and are often clavate, and the 

absence of secondary compounds. It is similar to 

the tempertae T. subtile, which, however, can be 

distinguished by the more distinctly emergent 

apothecia and ascospores that become brown at age. 


T. pseudosubtile. 

Thelotrema diplotrema is also similar, but differs by a less compact, sometimes roughened 

thallus surface, more distinctly emergent ascomata with a usually less distinctly free proper 

exciple and larger ascospores (up to 100 µm long with up to 22 loci) with thicker cell walls 

that lack a distinctly crenate surface. 

SPECIMENS EXAMINED – Australia, Queensland: Cape Tribulation Area: Myall Beach, Mangold 31 u (F); 

track to Cape Tribulation Beach, Mangold 32 t (F); 4.5 km on Buchanan [Creek] Rd., Hale 831723 (US). Km 45 

on Mt. Windsor Rd., NW of Mossman, Hale 831722 (US). Mt. Lewis Rd. 4 km N from Kennedy Hwy., W of 

Mossman, Hale 832737 (US). Manchans Beech, few km N of Cairns, K. & A. Kalb 20046 (hb. Kalb). W of Palm 

Cove, c. 25 km N of Cairns, K. & A. Kalb 19949 (hb. Kalb). Crystal Cascades, 5 km W of Cairns, Lumbsch & 

Mangold 19117 k (F). Atherton Tablelands: Lake Tinaroo, Downfall Creek Camping Area, Lumbsch & Mangold 

19125 j, h (F); 10 km S of Ravenshoe on Tully Falls Rd., Hale 830731 (US); 1-2 km N of Murray Falls, W of 

Kennedy, Hale 831342, 831648, 832533 (US). Eungella NP., along Broken River, Lumbsch & Mangold 19100 y


(F). Cape Hillsborough NP., NW of Mackay, Hale 831356, 831701 (US). Wooroi State Forest Park, W of 

Tewantin, Hale 830999, 832080 (US). Booloumba Creek SF., SW of Kenilworth, Hale 831367 (US). 6 km N of 

Jimna, Tibell 12797 (UPS). Bunya Mts.: On the rd. from the ridge to Maidenwell, 1.3 km NE of the intersection, 

Hafellner 19349 (GZU); Mt. Bunya, Shire picnic area on southern NP. boundary, S of Park, Hale 831411, 

831419, 832472, 832500 (US); Mt. Mowbullan, K. & A. Kalb 20254, 20268 (hb. Kalb). D'Anguilar Range NW 

of Brisbane, W of Mt. Glorious township, Hafellner 16955, 16957 (GZU). Mt. Mee SF., near Mt. Mee, N of 

Brisbane, Hale 58604 (US). Bennet Rd., Mt. Glorious, Rogers 8449 (BRI). Sankeys Shrub, Brisbane, Wilson s.n. 

(NSW-539342). Goodna [SW of Brisbane], Wilson s.n. (NSW-539346). Lamington NP.: Python Rock Track, 

Hale 832375 (US); Beechmont Range, Binna Burra, K. & A. Kalb 19896 (hb. Kalb). New South Wales: 

Nightcap Forest Drive, 1 km W of Minyon Falls, N of Lismore, Hale 832087, 832180, 832543 (US). Dorrigo 

NP., Never Never Picnic Area and Rosewood Creek Track, Mangold 24 b, d, f (F). Saltwater, E of Taree, Elix 

3997 (CANB). Sugar Creek Flora Reserve, Wallingat SF., 16 km SW of Forster, Streimann 44230 a (B, CANB). 

Bulahdelah District, Myall River SF., E of Stroud, Jarrah Rd., Kalb & Filson 18044, 18046, 18049 (hb. Kalb). 

Trail along bank of Mill Creek, 50 km NW of Sydney, K. & A. Kalb 34272 (hb. Kalb). Below Katoomba Falls, 

trail to Giant Stairway, Katoomba, Hale 58731 (US). Royal NP., S of Sydney, Bola Creek, E of Waterfall, K. & 

A. Kalb 21692, 21695, 21709 (hb. Kalb). Lord Howe Island: Goat House Cave, Elix 42175 (CANB), 42265 (B, 

CANB); Smoking Tree Ridge, Elix 42149 (B, CANB); Track from Smoking Tree Ridge to Rocky Run, Elix 

42429 (CANB); Track to Kims Lookout, Elix 42393 (CANB). 

Thelotrema rugatulum Nyl. 

Bull. Soc. Linn. Normand. 2(7): 168 (1873). Type: Andaman Islands, 1867, Kurz 57 (H-Nyl. 22495!lectotype, 

selected by Hale [1972 in hb.]). 


Thallus epi- to hypophloedal, (moderately) thin, up to c. 200 µm high, pale yellowish gray 

to more rarely pale grayish-green. Surface typical, dull to slightly shiny, usually smooth, 

sometimes the surface becoming partly eroded, then roughened to porous, rarely continuous 

to usually rugose and distinctly verrucose to slightly verrucose, ±fissured to areolate. Thallus 

cover variable, thallus predominantly covered by an ±incontinuous protocortex up to 25 µm 

thick, sometimes partly becoming distinctly conglutinated forming a true cortex of 

predominantly periclinal hyphae. Algal layer usually well developed, continuous to 

incontinuous, often interrupted by calcium oxalate crystals, calcium oxalate crystals very 

abundant, small to more often (very) large, often clustered. Vegetative propagules not seen. 

Ascomata variable, conspicuous to inconspicuous, moderately small to large, up to c. 1 mm in 

diam., ±roundish, peri- to apothecioid, erumpent to sessile, solitary to rarely marginally 

slightly fused, immersed to more rarely distinctly emergent, (verrucose-)hemispherical to 

more rarely (verrucose-)urceolate. Disc not visible from surface to rarely becoming partly 

visible, grayish, pruinose. Pores variable, small to wide to gaping, up to c. 800 µm in diam., 

predominantly smaller, usually up to 300 µm, roundish to roundish-irregular to irregular, 

proper exciple rarely entirely to more often only apically visible from surface, rarely entirely 

free to more often free only in the upper parts, usually off-white to whitish, often somewhat 

shrunken, predominantly entire to slightly split, in distinctly apothecioid specimen usually 

distinctly split, sometimes distinctly layered, incurved. Thalline rim margin ±roundish, small 

to gaping, predominantly entire to slightly split, sometimes slightly eroded, moderately thin to 

moderately thick, incurved, concolorous with thallus. Proper exciple apically free to rarely 

entirely free, moderately thin to moderately thick, with distinct, thick hyaline part internally, 

pale yellowish to yellowish-brown marginally, apically sometimes covered by grayish to 

rarely dark-gray granules, sometimes amyloid. Hymenium up to c. 250 µm high, noninspersed, 

moderately conglutinated, paraphyses parallel or slightly interwoven, unbranched, 

tips slightly thickened, lateral paraphyses present, often inconspicuous, up to c. 25 µm long, 

columellar structures absent. Epihymenium predominantly indistinct, hyaline and without 

granules, sometimes thin with small grayish granules and small crystals. Asci 1(-2)-spored,


tholus moderately thick to thin, not visible at maturity. Ascospores (very) large, densely 

eumuriform, cell walls and endospore (moderately) thin, often covered by a thin to 

moderately thick halo, hyaline, sometimes pale yellowish in old or decayed ascospores, 

faintly to more rarely distinctly amyloid, cylindrical to oblong-ellipsoid to broad-fusiform, 

with roundish to narrowed-roundish ends, loci roundish to angular, subglobular to irregular, 

transverse septae thin, distinct, usually very regular, 80-220 x 15-45 µm with multiple loci. 



Fig. 162. Thelotrema rugatulum: growth habit (A, B), ascomata (C), ascoma section (D), ascospore 

(E), ascospore detail (F) and ascospore showing amyloid reaction (G). A.: H-lectotype; B., D.-G.: 

Hale 831727; C.: Lumbsch & Mangold 19156 b. Bar= A: 1 mm; B: 0.8 mm; C: 0.7 mm; D: 150 

µm; E: 23 µm; F: 15 µm; G: 20 µm.



rugatulum was collected in Australia on tree bark 

in (sub)tropical rainforests, rarely in mangroves, in 

altitudes ranging from sea level to 900 m. It is rare 

but wide-spread, ranging from northern Queensland 

to southern New South Wales. This is the first 

report for Australia and New Zealand (see below). 

Previously only known from the type locality in the 

Andaman Islands. 

NOTES – Thelotrema rugatulum has abundant 

calcium oxalate crystals in the thallus and due to 

that a distinctly verruculose surface. It is further 

characterized by ascomata with an indistinctly free 

proper exciple, large, densely eumuriform, thinwalled, 

hyaline, ±amyloid ascospores and the 

absence of secondary compounds. A similar species 

is T. conveniens, see under that species for 

differences. Topeliopsis laceratula is another 

similar species, which can be distinguished by the 

distinctly greenish, smooth, constantly corticate thallus, strongly split and layered ascoma 

margins, a fused proper exciple and strongly amyloid, thick-walled ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Atherton Tablelands: Souita Falls, Lumbsch & Mangold 

19156 b (F); Just S of hwy., 13 km E of jct. Kennedy Hwy. and Palmerston Hwy., E of Ravenshoe, Hale 830847 

(US); 10 km S of Ravenshoe on Tully Falls Rd., Hale 832408 (US). Eungella NP., NP. side rd. nr. Peases 

Lookout, off Darymple rd., Hale 831645, 831727 (US). N side of Lake Baroon Pocket, W of Montville, 

Lumbsch & Mangold 19082 b (F). Mt. Mee SF., 6 km NW of Forestry Office, NW of Mt. Mee, Hale 831270, 

832351 (US). New South Wales: Southern bank of Hawkesbury River, along Singleton Rd. at Paddys Bight, K. 

& A. Kalb 34282 (hb. Kalb). New Zealand, South Island, Nelson, 25.10.2005, Polly s.n. (WELT). 

Thelotrema saxatile C. Knight 

Trans. and Proc. New Zealand Inst. 8: 327 (1876). Leptotrema saxatile (Knight) Müll. Arg., Bull. Soc. Bot. 

Belgique 31: 35 (1892). Leptotrema monosporum var. saxatile (Knight) Müll. Arg., Bull. Herb. Boissier 2(1): 75 

(1894). Type: New Zealand, (?Wellington), 01.Feb.1882, C. Knight s.n. (WELT-hb. Knight[36: 12]-lectotype, 

selected by Galloway [1985: 577]; BM!-isolectotype). 

Ascidium manosporum Knight, Trans. New Zealand Inst. 15: 355 (1883). Leptotrema manosporum (Knight) 

Müll. Arg., Bull. Herb. Boissier 2(1): 75 (1894). Thelotrema manosporum (Knight) Hellb., Bihang till Kgl. 

Svensk. Vetensk.-Akad. Handl. vol. 21, 3(13): 78 (1896) [the epithet is corrected to A. monosporum on the type 

annotation and Galloway (1985) uses this name, however, it was published as A. manosporum]. Type: New 

Zealand, (?Wellington), C. Knight s.n. (WELT-hb. Knight[37A: 1]!-lectotype, selected by Galloway [1985: 575 

as A. monosporum]). 

Thelotrema monosporoides Nyl., Lich. Nov. Zealand p. 76 (1888). Thelotrema monosporum Kremp. nom. 

illeg., Verhandl. zool.-bot. Gesellsch. Wien 26: 453 (1876) [non Thelotrema monosporum Nyl.]. Leptotrema 

monosporoides (Nyl.) Müll. Arg., Bull. Herb. Boissier 2(1): 75 (1894). Ocellularia monosporoides (Nyl.) Hale, 

Mycotaxon 11: 137 (1980). Type: New Zealand, (?Wellington), C. Knight s.n. (H-Nyl. 22708!-lectotype, 

selected by selected by Hale [1981: 314]; G-isolectotype). 

Thelotrema monospermum Harris, Some Florida Lichens: 99 (1990). Type: U.S.A., Florida, Liberty County, 

Harris 11399 (NY!-holotype). 



T. rugatulum.


Fig. 164. Thelotrema saxatile: growth habit (A, C), ascomata (B, D), ascoma section (E), young, 

mature and over-mature ascospores (F-H) and ascospore detail (I). A., E.-G.: BM-isolectotype; B.: 

WELT-lectotype of A. manosporum; C.: NY-lectotype of T. monospermum; D.: Mangold 32 v. 

Bar= A: 2 mm; B: 1 mm; C: 2.5 mm; D: 0.5 mm; E: 175 µm; F: 40 µm; G: 25 µm; H: 18 µm; I: 

12.5 µm. 

Thallus variable, corticolous to rarely saxicolous, epi- to predominantly hypophloedal, epito 

somewhat endolithic in saxicolous specimen, moderately thin to moderately thick in 

saxicolous specimen, up to c. 300 µm high, pale yellowish-brown to pale grayish-green or 

whitish-gray. Surface variable, dull to somewhat glittering, predominantly smooth to rarely 

roughened, continuous to verrucose or verruculose, unfissured to fissured. True cortex usually 

absent, thallus predominantly covered by a continuous to incontinuous protocortex up to


25 µm thick, very rarely becoming distinctly conglutinated forming a thin true cortex of 

periclinal hyphae. Algal layer variable, well to poorly developed, continuous to incontinuous, 

calcium oxalate usually abundant, sometimes sparse, small to large, scattered or clustered. 

Vegetative propagules not seen. Ascomata variable, conspicuous to inconspicuous, 

moderately small to large, up to 800 µm in diam., ±roundish, peri- to predominantly 

apothecioid, usually sessile, solitary to marginally fused, immersed to more often ±distinctly 

emergent, (irregular-)hemispherical to more rarely urceolate to very rarely irregularsubglobose. 

Disc often becoming partly visible from surface, pale grayish to gray, moderately 

to strongly pruinose. Pores variable, small to wide to gaping, up to c. 600 µm, roundish to 

more often roundish-irregular to irregular, proper exciple entirely to apically visible from 

surface, entirely free, usually off-white to whitish, often shrunken, predominantly incurved to 

more rarely erect. Thalline rim margin thin to more often moderately thin to thick, roundish to 

irregular, small to gaping, entire to ±distinctly split to somewhat lacerate or eroded, incurved 

to sometimes erect, concolorous with thallus to brownish or more rarely brighter than thallus. 

Proper exciple ±free, moderately thin to moderately thick, hyaline to pale yellowish 

internally, yellowish-brown marginally, apically often brownish to dark-brown and covered 

by dark-gray granules, sometimes amyloid at the base. Hymenium up to c. 250 µm high, noninspersed, 

moderately conglutinated, paraphyses parallel or slightly interwoven, unbranched, 

tips moderately to distinctly thickened, lateral paraphyses present, often inconspicuous, up to 

c. 20 µm long, columellar structures absent. Epihymenium variably thick, hyaline, usually 

with fine to coarse grayish granules and small crystals. Asci 1-2-spored, tholus usually thin or 

not visible. Ascospores (very) large, densely eumuriform, in immature stages often with 

distinctly thickened cell walls, at maturity cell walls and endospore predominantly 

(moderately) thin, non-halonate, brown at early maturity, in very large ascospores tips often 

remaining unpigmented, non-amyloid, cylindrical to oblong-ellipsoid, with roundish to 

narrowed-roundish ends, loci predominantly roundish to slightly angular, subglobular to 

somewhat oblong, transverse septae thin, distinct throughout development, regular, 70- 

180(200) x 15-40 µm with c. 24-46 x 2-10 loci. Pycnidia not seen. 



saxatile was collected in Australia on tree bark in 


sea level to 1100 m. It is a rare but wide-spread 

species occurring in northern and north-central 

Queensland and on Lord Howe Island (New South 

Wales). This is the first report for Australia. The 

distribution is not entirely clear, due to difficulties 

in the distinction from T. monosporum (see there). I 

have seen material from the U.S.A. and New 

Zealand 

N OTES – Thelotrema saxatile is a morphologically 

variable species. It is characterized by the 

often dark, free exciple, single- to more rarely 

double-spored asci with large, densely eumuriform, 

brown, thick-walled immature and thin-walled 

mature, non-amyloid, predominantly cylindrical, 

never distinctly fusiform ascospores and the 

absence of secondary compounds. It has been 


T. saxatile.


considered conspecific with T. monosporum ever since Salisbury (1972). The circumscription 

of this similar taxon was always unclear and thus has caused many confusion. Supported by 

molecular data (Lumbsch & al., 2007) it could be shown that the taxa formerly merged under 

the name T. monosporum s. lat. can be assigned to three distinct lineages, viz. T. monosporum 

s. str., T. lepadodes and T. saxatile, which can be set apart by their ascospore morphologies. 

Thelotrema saxatile differs from T. monosporum by single- to more rarely 2-spored asci (up 

to four spores per ascus in T. monosporum) with distinctly larger, cylindrical ascospores and a 

thicker, more compact, often distinctly verrucose or verruculose thallus. Similar and probably 

synonymous to T. saxatile is T. macrosporum from Europe (Purvis & al., 1995). However, the 

type was not available for study. Another similar Australian Thelotrema that lacks chemistry 

and has large, muriform, brownish ascospores is T. conveniens. This species can be readily 

distinguished, however, by distinctly amyloid ascospores that become pigmented. 

SPECIMENS EXAMINED – Australia, Queensland: Iron Range NP., York Peninsula, 31 km from western 

boundary on track to Portland Rds., Hale 832628 (US). Cape Tribulation area, track to Cape Tribulation Beach, 

Mangold 32 h, j, o, p, v (F). Mt. Windsor, 5 km W of new Forestry Camp, NW of Mossman, Hale 831953 (US). 

Eungella NP.: Along Broken River, Lumbsch & Mangold 19104 a (F); Finch Hatton Gorge, Lumbsch & 

Mangold 19113 l (F). New South Wales: Lord Howe Island, track to little Island, near Salmon Beach, Elix 

32664 a (B). 

Thelotrema saxicola (Vain.) Salisb. 

Lichenol. 5: 269 (1972). Bas.: Thelotrema lepadinum ssp. saxicola Vain., Étud. Lich. Brésil 3: 77 (1890). 

Leptotrema saxicola (Vain.) Redgr., Arkiv för Bot. 28A, 8: 102 (1936). Type: Brazil, Minas Gerais, Vainio s.n. 

(Lich. Brasil. exsicc. 1173) (TUR-Vain. 26784!-holotype). 


Thallus saxi- or corticolous, epilithic or epi- to hypophloedal, moderately thick, up to c. 

400 µm high, grayish-green to pale yellowish. Surface dull to slightly shiny, smooth, rarely 

continuous to usually distinctly to strongly verrucose or verruculose, distinctly fissured to 

somewhat areolate. Thallus cover variable, predominantly covered by a continuous to 

incontinuous protocortex, sometimes becoming distinctly conglutinated, forming a true cortex 

of periclinal hyphae, up to c. 20 µm thick. Algal layer moderately well developed, 

±continuous, calcium oxalate crystals sparse to moderately frequent, small to large and 

usually scattered. Vegetative propagules not seen. Ascomata predominantly inconspicuous, 

(moderately) large, up to c. 600 µm in diam., roundish, solitary to marginally fused, sessile, 

perithecioid, immersed to emergent, then predominantly (verrucose-) hemispherical or 

somewhat cylindrical (deceased, sterile ascomata sometimes resembling conspicuous, 

apothecioid ascomata). Disc not visible from surface. Pores predominantly very small, up to 

c. 50(100) µm in diam., roundish to slightly irregular, entire to slightly split, proper exciple 

margin sometimes visible from surface, fused to rarely slightly detached, whitish-translucent 

or pale to dark gray, flush with thalline rim margin or somewhat sunken. Thalline rim margin 

(moderately) thick, roundish, predominantly entire, concolorous with thallus or brighter, flush 

with thalline rim or slightly depressed, thalline rim incurved, with same surface as thallus. 

Proper exciple predominantly fused, moderately thin, hyaline internally, pale yellowish to 

yellowish-brown marginally, apically often dark-brown to slightly carbonized, in corticolous 

specimen often with an indistinct marginal layer of substrate material, sometimes slightly 

amyloid (reddish) to distinctly amyloid (purple) at the base. Hymenium up to c. 350 µm high, 

inspersed, distinctly conglutinated, paraphyses thin, ±interwoven, unbranched, straight to


Fig. 166. Thelotrema saxicola: growth habit (A), ascomata (B), ascospore (C) and ascospore detail 

(D). A., B., D.: TUR-holotype; C.: Hale 831753. Bar= A: 1.2 mm; B: 0.3 mm; C: 30 µm; D: 8 µm. 

somewhat bent, tips unthickened, lateral paraphyses present variably conspicuous, up to 

40 µm long, columellar structures absent. Epihymenium indistinct, hyaline, without granules 

or crystals. Asci 1-spored, tholus thin, not visible at maturity. Ascospores predominantly very 

large, densely eumuriform, cell walls thin to slightly thickened, endospore absent, nonhalonate, 

hyaline, becoming yellowish to brownish in late maturity, non-amyloid to usually 

weakly amyloid in older stages, predominantly oblong to more rarely somewhat fusiform with 

roundish to narrowed-roundish ends, loci small, 

roundish to slightly angular, subglobose to 

irregular, transverse septae thin, distinct in younger 

stages, becoming indistinct with age, ±regular, 150- 

250 x 30-50 µm with multiple loci. Pycnidia not 

seen. 

CHEMISTRY – Thallus K+ yellowish, C-, PD+ 

yellow; containing psoromic (major), 2'-0-demethylpsoromic 

and subpsoromic (traces) acids. 


saxicola grows in Australia on tree bark in (sub-)tropical 

rainforests. It is rare and disjunct in 

northern Queensland and northern New South 

Wales. This is the first report for Australia. 

Previously only known from Brazil. 

N OTES – This species is recognized by 

perithecioid ascomata with an inspersed hymenium, 


T. saxicola.


lateral paraphyses and dark exciple, large, eumuriform, moderately pigmented asco-spores 

and the presence of the psoromic acid chemosydrome. Thelotrema oleosum is similar but 

differs in the absence of secondary metabolites. Further differences include more open-pored 

(pores up to 150 µm in diam.) ascomata with more distinctly detached proper exciple, asci 

with thickened lateral walls and ascospores that generally show a stronger amyloid reaction. 

Also similar is T. porinaceum, which can be readily distinguished by a thallus with an 

indistinct cortex, ascomata with more distinctly detached proper exciple, unpigmented 

ascospores, and the occurrence of norstictic acid. 

SPECIMENS EXAMINED – Australia, Queensland: Km 45 on Mt. Windsor Rd., NW of Mossman, Hale 832753 

(US). New South Wales: Big Shrub Flora Reserve, Night Cap Forest Drive, W of Mullumbimby, Hale 831402, 

831680 (US). Cambridge Plateau Forest Drive, 1-3 km N of picnic area, Richmond Range SF., 30 km W of 

Casino, Hale 831627 (US). 

Thelotrema subadjectum Mangold spec. nov. ined. 

Type: Australia, Queensland, Mt. Bunya, Shire picnic area to National Park boundary, south of park, Hale 

831440 (US-holotype). 

ETYMOLOGY – The epithet refers to the similarities to T. adjectum. 


Thallus predominantly hypophloedal, rarely a very thin epiphloedal part developing, up to 

c. 100 µm high, pale grayish to pale grayish-green. Surface dull to somewhat glittering, 

slightly pruinose to roughened, usually due to protuberate substrate surface, continuous, 

slightly fissured to unfissured. Cortex structures absent. Algal layer poorly to moderately well 

developed, ±continuous, calcium oxalate sparse, often lacking in major parts, ±small, often 

clustered. Vegetative propagules not seen. Ascomata inconspicuous, moderately small, up to 

c. 600 µm in diam., roundish, apothecioid, solitary to rarely marginally fused, usually 

distinctly immersed. Disc not visible from surface to very rarely somewhat visible, pale 

grayish to whitish, pruinose. Pores small, up to c. 150 µm in diam., roundish to roundishirregular 

to irregular, entire to slightly split, apical proper exciple visible from surface, 

whitish, incurved, more rarely slightly shrunken. Thalline rim margin moderately thick, 

roundish, entire to slightly split, sometimes slightly funnel-shaped, concolorous with thallus 

to whitish. Proper exciple becoming free with age, thick, hyaline internally, pale brownish to 

grayish marginally, often with substrate particles incorporated, apically often covered by 

grayish granules, usually amyloid at the base and the marginal parts of the subhymenium. 


parallel to slightly interwoven, unbranched, tips slightly thickened, lateral paraphyses present, 

conspicuous, up to c. 25 µm long, columellar structures absent. Epihymenium thin, hyaline, 

with fine to coarse grayish granules. Asci 8-spored, tholus thick, thin when mature. 

Ascospores (moderately) small, (sub-)muriform, cell walls moderately thick, endospore thick, 

distinctly halonate, hyaline, faintly amyloid, predominantly oblong to somewhat fusi- or 

claviform, with roundish to narrowed-roundish ends, loci roundish to slightly angular, 

subglobose or irregular to more often ±oblong, immature ascospores with large, conical end 

cells, otherwise hemispherical to conform with other loci, transverse septae (moderately) 

thick, distinct and regular, 25-35 x 8-10 µm with 10-14 x 1-4 loci. Pycnidia not seen. 

CHEMISTRY – Thallus K-, C-, PD-; no secondary compounds detectable by TLC.


Fig. 168. Thelotrema subadjectum: growth habit (A), ascomata (B) and ascospores (C, D). A.-D.: 

US-holotype. Bar= A: 2 mm; B: 0.7 mm; C: 7 µm; D: 5 µm. 

ECOLOGY AND DISTRIBUTION – Thelotrema subadjectum 

grows on tree bark in (sub)tropical 

rainforests. It is rare and currently only known in 

northern and southern Queensland. 

NOTES – This new taxon is characterized by a 

hypophloedal, ecorticate thallus, immersed, small 

ascomata with a free bright proper exciple, 

moderately small, muriform, hyaline, weakly 

amyloid ascospores with thickened endospore and 

transverse septae, and the absence of secondary 

metabolites. It can be distinguished from the similar 

T. adjectum by a somewhat thinner thallus, never 

emergent ascomata with distinctly free and visible 

proper exciple, unlayered thalline rims and smaller 

ascospores (up to 80 µm long in T. adjectum). Also 

similar are T. polythecium and T. cyphelloides. 

Thelotrema polythecium is readily distinguished by 


T. subadjectum.


ascomata with distinctly split margins and fused proper exciple and smaller, non-amyloid 

ascospores (up to 25 µm long with up to 9 x 4 loci). Thelotrema cyphelloides differs by 

having smaller ascospores (up to 27 µm long with up to 9 x 4 loci) and the presence of the 

stictic acid chemosyndrome. 

SPECIMENS EXAMINED – Australia, Queensland, 5 km NW of Babinda, near bridge crossing Russell River, 

Hale 831112 (US). 

Thelotrema subtile Tuck. 

Amer. J. Arts and Sci., ser. 2, 25: 426 (1858). Ocellularia subtilis (Tuck.) Riddle, Mycologia 15: 79 (1923). 

Type: U.S.A., Vermont, Brattleboro, 1851, Frost 150 pr. p. (FH-Tuck.!-lectotype, newly selected here 13 ; FH- 

Tuck.!- [pr. p. as Frost 96], NY!-isolectotypes). 

Ocellularia jugalis Müll.Arg., Bull. Herb. Boissier 3: 313 (1895). Type: Australia, Queensland, Sankeys 

Shrub [Brisbane], 1893, Shirley 1836 (G!-holotype, BRI-'Shirley Book', p. 21, n. 35 [BRI-AQ721220]!-isotype). 



brown to pale grayish or tannish-gray. Surface dull to slightly shiny, smooth, continuous to 

more often ±distinctly verrucose to verruculose slightly fissured to unfissured. Thallus cover 

quite variable, predominantly an incontinuous protocortex present, up to c. 25 µm thick, 

sometimes becoming distinctly conglutinated forming a true cortex of periclinal hyphae. 

Algal layer variable, well to poorly developed, continuous to incontinuous, calcium oxalate 

abundant, small to large, scattered or clustered. Vegetative propagules not seen. Ascomata 

conspicuous, predominantly small to more rarely moderately large, up to c. 600 µm in diam., 

roundish to somewhat irregular, apothecioid, sessile, solitary to marginally or rarely entirely 

fused, predominantly ±distinctly emergent, hemispherical to urceolate or subglobose, with 

same surface as thallus or more distinctly verruculose. Disc usually becoming partly visible 

from surface in older ascomata, (dark)gray, ±distinctly pruinose. Pores small to moderately 

wide to rarely gaping, up to c. 400 µm in diam., roundish to irregular, entire to slightly split, 

proper exciple apically to more often entirely visible from surface, apically bright to brownish 

towards the base, sometimes shrunken, incurved to somewhat erect. Thalline rim margin thin 

to moderately thick, usually becoming wide to gaping, entire to split to more rarely somewhat 

eroded, roundish to irregular-roundish, predominantly incurved and concolorous with thallus. 

Proper exciple free, moderately thick, hyaline to predominantly pale yellowish internally, 

yellowish- to grayish-brown marginally, apically often dark-brown, sometimes with substrate 

inclusions, rarely amyloid at the base. Hymenium up to c. 150 µm high, non-inspersed, 

distinctly conglutinated, paraphyses parallel to slightly interwoven, unbranched to sometimes 

slightly branched, tips irregular, moderately thickened, lateral paraphyses present, usually 


thick, hyaline to sometimes brownish with age, usually with fine grayish-brown granules and 

small crystals. Asci 4-8-spored, tholus (moderately) thick, thin when mature. Ascospores 

13 Salisbury’s (1972) type-selection (Frost, Reliquiae Tuckermanianae no. 140) consists of four collections (on 

bark of Acer, Betula and Fraxinus), that are intermixed and shared in two capsules. One capsule is labeled "On 

Acer, Frost(96), 1836" (FH-213376), the other "Frost(150), 1851" (FH-213377). Obviously Tuckerman selected 

the latter sample (FH-213377) as the 'illustrative material' since it consists of a sheet with several pieces of two 

specimens glued on, and has descriptive notes and a drawing of an ascospore. It is also labeled as lectotype by 

Hale 1972 (unpublished). The specimen on the left (on Fraxinus) is the here selected lectotype. The specimen on 

the right (on Betula, determined as T. suecicum by Maass) is also T. subtile. Several pieces of this collection are 

also included in the first mentioned capsule (FH-213376) together with a specimen of T. subtile on Acer and a 

collection of T. suecicum (on Betula). A duplicate of the type collection is also deposited in NY.


moderately small, transversely septate, cell walls (moderately) thick, often with distinctly 

crenate surface, with thin halo, hyaline in young and mature stages, becoming distinctly 

(pale)brownish in over-mature or decayed ascospores, faintly to moderately amyloid, 

(oblong)fusiform to more often clavate, ends roundish to acute, loci roundish to slightly 

angular, rarely subglobose to more often lentiform, end cells hemispherical to conical, septae 

(moderately) thick, regular, 30-50 x 7-10 µm with 8-16 loci. Pycnidia not seen. 


Fig. 170. Thelotrema subtile: growth habit (A. B), ascoma section (C), ascospores (D, E) and 

ascospore showing amyloid reaction (F). A., C., E.: FH-lectotype; B.: G-holotype of O. jugalis; 

D., F.: Mangold 3 e. Bar= A: 0.75 mm; B: 0.6 mm; C: 150 µm; D, F: 6 µm; E: 8 µm.



subtile grows on bark in cool-temperate rainforests 

and wet sclerophyll forests, rarely in sub-tropics 

(see notes), in altitudes ranging from 250 to 500 m. 

It is rare in Australia, occurring in southern 

Queensland (see notes), southern Victoria and 

north-eastern Tasmania. It is also known from 

Hawaii (Salisbury, 1975), the Neotropics, Africa 

(Frisch, 2006), Japan (Matsumoto, 2000), 

Philippines (Salisbury, 1975), Java (ibid.), New 

Caledonia (ibid.) and New Zealand (ibid.), 

indicating a pantemperate to pan(sub)tropical distribution 

(see also below). 

NOTES – This species is characterized by a thin 

thallus, predominantly ±distinctly emergent, small 

apothecia with free proper exciple, moderately 

small, transversely septate, weak to moderately 

amyloid ascospores with a crenate surface, that are 

hyaline at maturity but become distinctly brown, 


T. subtile. 

and the absence of secondary compounds. Several similar species are known, however, T. 

subtile is distinguished from all members of Thelotrema with similar, hyaline, (moderately) 

small, transversely septate ascospores that lack secondary compounds by the pigmentation of 

the ascospores in the terminal phases of their development. The most similar T. pseudosubtile 

additionally differs by usually immersed or only slightly raised ascomata with less distinctly 

free proper exciple. Thelotrema defossum differs by distinctly smaller ascospores 

(predominantly up to 30 µm with up to 11 loci), for additional differences to T. diplotrema 

and T. suecicum see under these species. Two further similar Australian species, T. 

circumscriptum and T. bicavatum, differ by the presence of secondary compounds, see there 

for additional differences. Out of the here examined specimen only Shirley’s collection from 

Brisbane (the type of O. jugalis) and one collection from South Carolina are originated in 

subtropical or warm-temperate climates, the type collection as well as all other Australian 

collections were made in cool-temperate areas. Thus, it is assumed that many of the 

specimens reported from the tropics are likely to be misidentified and probably are in fact one 

of the other similar taxa occurring in the tropics. Salisbury (1975) reports T. subtile based on 

a Wilson collection from New South Wales (also listed in Frisch, 2006), however, the two 

specimen in H are O. wirthii. A careful re-examination of the herbarium material is needed 

and the distribution of T. subtile is only tentative. 

SPECIMENS EXAMINED – Australia, Victoria: Warburton, Dec. 1885, Wilson '513' (NSW-539393). Ferntree 

Gully: 28. Jan.1892, Wilson s.n. (NSW-539396); 23. Dec.1899, Bastow s.n. (MEL-724546). Ottway Ranges, 10 

km N of Apollo Bay, Mangold 3 e, j (F). Ottway NP., 10 km W of Apollo Bay, Mangold 1 d (F). Tasmania: St. 

Marys, Wilson s.n. (NSW-539323). U.S.A.: Vermont, Brattleboro: '1851, Frost 150' (with lectotype, on Betula) 

(FH-213377 pr.p.); '1836, Frost 96' (with isolectotype, on Acer) (FH-213376 pr.p., WIS [as Reliq.Tuck.140]). 

South Carolina, Ravenel s.n. (Reliq.Tuck.141) (NY, WIS).


Thelotrema suecicum (Magn.) James 

Lichenologist 9: 186 (1977). Bas.: Ocellularia suecica Magn., Bot. Not. 1937: 125 (1937). Type: Sweden, 

Bohuslän, 22. Jun.1936, Magnusson s.n. (Lich. sel. scandin. exs. n. 230) (BM!-lectotype, selected by James 

[1977: 186]; C!-, F!-isolectotypes). 

Ocellularia bonplandiae var. obliterata Müll. Arg., Bull. Herb. Boissier 1: 54 (1893). Type: Australia, 

Victoria, Warburton, Wilson 513 (G!-holotype, NSW!-isotype). 


Fig. 172. Thelotrema suecicum: growth habit (A, B), ascomata (C) and ascospores (D, E). A.: 

Mangold 3 p; B.: G-lectotype of O. bonplandiae var. obliterata; C.: C-isolectotype; D.: Hafellner 

16634; E.: Kantvilas 670/90. Bar= A: 1.2 mm; B: 2 mm; C: 0.6 µm; D: 10 µm; E: 8 µm. 


gray to pale grayish-green or olive. Surface dull to slightly shiny, smooth, continuous to 

±distinctly verruculose, slightly fissured to unfissured. Thallus cover variable, predominantly 

an incontinuous protocortex present, up to c. 20 µm thick, sometimes becoming distinctly 

conglutinated forming a thin true cortex of periclinal hyphae Algal layer usually ±well 

developed and continuous, often becoming somewhat incontinuous due to calcium oxalate 

crystal inclusions, calcium oxalate crystals abundant, small to large, scattered to clustered. 

Vegetative propagules not seen. Ascomata conspicuous, predominantly small to more rarely 

moderately large, up to c. 700 µm in diam., roundish, apothecioid, sessile, solitary to 

sometimes marginally fused, rarely immersed to predominantly distinctly emergent, 

hemispherical to more rarely urceolate or subglobose, with same surface as thallus. Disc often 

becoming partly visible from surface, grayish, pruinose. Pores small to moderately wide to 

rarely gaping, up to c. 400 µm in diam., roundish to irregular, entire to split, apical to 

sometimes entire proper exciple visible from surface, often shrunken, apically bright, pale


brownish towards the base, incurved to rarely somewhat erect. Thalline rim margin thin to 

moderately thick, entire to split to somewhat eroded, ±roundish, usually becoming wide to 

gaping, incurved to slightly erect, concolorous with thallus or yellowish to pale orange. 

Proper exciple free, (moderately) thin, hyaline to pale yellowish internally, yellowish to 

brownish marginally, apically covered by grayish granules, sometimes slightly amyloid at the 

base. Hymenium up to c. 130 µm high, non-inspersed, weakly conglutinated, paraphyses 

parallel to slightly interwoven, unbranched, tips moderately thickened, lateral paraphyses 

present, usually inconspicuous, up to c. 25 µm long, columellar structures absent. 

Epihymenium (moderately) thin, hyaline, with grayish granules and sometimes small crystals. 

Asci 8-spored, tholus (moderately) thick, thin at maturity. Ascospores typical, (moderately) 

small, transversely septate, cell walls thick from very young stages on to very thick at 

maturity, moderately thin to moderately thick halo, hyaline, non-amyloid to usually only very 

faintly amyloid, predominantly ellipsoid to broad-fusiform to more rarely broad-clavate, ends 

narrowed-roundish to more rarely subacute, loci roundish to angular, subglobose to cuboid or 

irregular, latitudinal as well as longitudinal elongate , end cells hemispherical to conical, 

septae (moderately) thick, ±irregular, 20-40(60) x 8-15 µm with 6-12(14) loci. Pycnidia not 

seen. 


ECOLOGY AND DISTRIBUTION – The species 

grows on tree bark predominantly in sub-tropical to 

cool-temperate rainforests, more rarely in wet 

sclerophyll forests and very rarely in tropical 

highland rainforest, in altitudes ranging from 20 to 

1400 m. This common and wide-spread species 

occurs from northern Queensland to southern 

Victoria, and Tasmania. It is also known from 

North and South America (Purvis & al., 1995), 

Europe and New Zealand (Lumbsch & al., 2008) 

indicating a subcosmopolitan distribution. 

NOTES – Thelotrema suecicum is characterized a 

thin thallus, predominantly distinctly emergent, 

small apothecia with free proper exciple, 

moderately small, hyaline, transversely septate 

ascospores with thick cell walls, even in immature 

stages. The ascospores further show no or a very 

faint amyloid reaction, the loci are divided by thick 

septae, distinctly irregular formed and often 


T. suecicum. 

longitudinal elongated. The ascospore sizes of the type and in most of the Australian 

specimens do not extend lengths of ±30 µm, however, in some collections, predominantly 

from Tasmania, several larger ascospores were found, in some cases reaching up to 60 µm 

with up to 14 loci. Two similar species are T. defossum and T. subtile. The (sub)tropical T. 

defossum can be readily distinguished by the moderately thin- to moderately thick-walled 

ascospores with regularly formed loci and thin septae. Thelotrema subtile is more difficult to 

distinguish, but the ascospores are more distinct amyloid, have less thickened cell walls that 

often show a crenate surface, larger loci and a brown pigmentation in deceased ascospores. 

SPECIMENS EXAMINED – Australia, Queensland: Atherton Tablelands, Millaa Millaa near the falls, M. & A. 

Aptroot 46167 h (ABL). Brisbane, 1891, Bailey s.n. (G-10194/41). New South Wales: Mt. Warning NP., track 

from summit to parking lot, Mangold 19 s (F). Border Ranges NP.: McPherson Range, NE of Gradys Creek,


Hafellner 16634 (GZU); Tweed Range, NE of Wiangaree, The Oinnacle, Hafellner 19171 (GZU). Track to 

Wrights Lookout, New England NP., 72 km E of Armidale, Elix 33931 (CANB). Berrico Rd., Chichester SF., 21 

km SW of Gloucester, Elix 25022 (CANB). Mt. William, Barrington Tops NP., 30. Jun.1888, Kantvilas s.n. 

(NSW-221836). Mt. Wilson: Ewers 3089 (CANB); Mt. Irvine Rd., 25 km NNE of Katoomba, Streimann 31589 

(CANB). Katoomba, Sept. 1889, Wilson s.n. (H). Monga SF./Monga NP.: 27 km SE of Braidwood, Mangold 11 

b (F); 5 km S of Monga, along Mongarlowe River, Kalb & Elix 21781, 21861 (hb. Kalb); ['Mongala River 

crossing Mongala State Forest'], 02. May 1986, Everett s.n. (NSW-539331). Brown Mt., c. 30 km SE of 

Nimmitabel, Kalb & Elix 30435 (hb. Kalb). Victoria: Ellery Camp, 30 km SSW of Bendoc, Elix 21441 (CANB). 

Gippsland ['Gypsland'], 1889, C. French s.n. (G-10194/48). Bonang rd., NE of Orbost, Mangold 8 d, e (F). 

Wilsons Promontory NP., on sealers cove hiking track, Mangold 6 l (F). Cumberland Falls, c. 19 km E of 

Marysville, Filson 1035395 (MEL). Eastern Highlands, Black Spur: 20. Mar.1885, Wilson s.n. (MEL-1063643, 

NSW-539397, -539419); 1888, Wilson s.n. (NSW-539391, -539397); Jan. 1890, Wilson s.n. (NSW-153541). 

River Watts [Yarra Ranges], 1891, Martin s.n. (NSW-539414). Fern Tree Gully [E of Melbourne]: 28. Jan.1892, 

Wilson s.n. (NSW-539396); 23. Dec.1899, Bastow s.n. (MEL-26165). Dandy Hills, Melbourne ['Danndy Hills'], 

Jan. 1892, Wilson s.n. (NSW-603861). Mt. Macedon [NW of Melbourne], Dec. 1891, Wilson s.n. (NSW- 

603846). Ottway Ranges, 10-14 km N of Apollo Bay, Mangold 3 p, 5 f (F). Tasmania: 3.5 km W of Luina, 

Kantvilas 332/89 (HO). Anthony Rd., Kantvilas 185/91, 434/91, 222/99, 189/93 (HO). Cradle Mtn. Lake St 

Clair NP., Pelion Plains, 1 km SW of Pelion Hut, Kantvilas 210/92 (HO). 3 km S of Teepookana, Kantvilas 

666/90, 670/90 (HO). Mt. Wellington, Wilson s.n. (NSW-539392). Warra Creek, 19. Jun.1996, Kantvilas s.n. 

(HO). W of Tahune Bridge, "Big Coupe - understorey island near 418 m.peg", in the Warra SST: Kantvilas 

222/99 (HO); 05. May 1998, Kantvilas s.n. (HO). Norway, Hordaland, Gaarder 4365, 4366 (BG). U.S.A, 

Vermont, Brattleboro 1836, Frost 96 (FH-213376 pr.p. [with isolectotype of T. subtile]). 

Thelotrema thesaurum Mangold spec. nov. ined. 

Type: Australia, Queensland, Dawson logging area, 24 km S of Koombooloomba turnoff, WSW of Tully, 

Hale 832638 (US-holotype). 

ETYMOLOGY – From lat. thesaurus (= treasure , store, hoard), the epithet refers to the 

ascomata of the new species with thick thalline rim and conspicuously large crystal 

aggregates. 


Thallus epi- to hypophloedal, thin to moderately thick, up to c. 300 µm thick, yellowishbrown 

to pale olive. Surface waxy, smooth, ±verrucose, fissured. True cortex present, 

yellowish, consisting of irregular hyphae, up to c.50 µm thick. Algal layer continuous and 

well developed, calcium oxalate crystals abundant, in particular in ascocarp area, (very) 

large, solitary to more rarely in clusters. Vegetative propagules not seen. Ascomata 

conspicuous, (very) large, up to c. 2 mm in diam., roundish, peri- to indistinctly apothecioid, 

sessile, solitary to fused, emergent, subglobose to urceolate. Discs not visible from surface. 

Pores (moderately) small, up to c. 300 µm in diam., roundish to slightly irregular, apical to 

proper exciple visible from surface, free, predominantly entire to slightly split, whitish to offwhite, 

predominantly incurved, usually ±shrunken. Thalline rim margin thick to very thick, 

roundish, entire to slightly split, in older stages usually becoming eroded, whitish to off-white 

and distinctly pruinose when eroded, thalline rim predominantly concolorous with thallus, 

with same surface as thallus, often distinctly fissured, incurved. Proper exciple free, 

moderately thin to moderately thick, hyaline to pale yellowish internally, yellowish-brown 

marginally, apically covered with grayish granules, non-amyloid to sometimes amyloid at the 


±bent, ±interwoven, tips slightly thickened and somewhat irregular, lateral paraphyses 

present, conspicuous, up to c. 30 µm long, columellar structures absent. Epihymenium 

moderately thick, yellowish-brown, with grayish to brownish granules. Asci (4)6-8-spored, 

tholus thick, thin when mature. Ascospores (moderately) large, densely eumuriform, cell 

walls and endospore (moderately) thin, non-halonate, hyaline to rarely pale yellowish with


age, non-amyloid to faintly amyloid, oblong to ellipsoid with roundish to narrowed-roundish 

ends, loci roundish to angular, subglobose to irregular, transverse septae thin, regular to 

slightly irregular, becoming indistinct at late maturity, 60-120(130) x 20-30 µm with multiple 


Fig. 174. Thelotrema thesaurum: growth habit (A, C), ascomata (B), ascoma section (D) and 

ascospores (E, F). A.: Streimann 45596; B.-F.: US-holotype. Bar= A: 2.5 mm; B: 0.75 mm; C: 1.5 

mm; D: 125 µm; E: 35 µm; F: 30 µm. 


constictic (majors), cryptostictic, α-acetylconstictic, hypoconstictic and hypostictic (traces) 

acids. 

ECOLOGY AND D ISTRIBUTION – Thelotrema thesaurum grows on bark in tropical 

rainforests, predominantly in highlands, in altitudes ranging from 100 to 1100 m. It is rare, 

known only from northern Queensland.


N OTES – Characteristic for this taxon are the 

large, in older stages apically eroded and brightpruinose 

ascomata with thick thalline rims with 

abundant, large calcium oxalate crystal inclusions. 

It is further characterized by a dark, waxy, 

corticate, ±verrucose thallus, large, eumuriform, 

thin-walled, hyaline, non- to faintly amyloid 

ascospores and the presence of the stictic acid 

chemosyndrome. It is similar to T. leucophthalmum 

and T. cupulare, both distinguished by 

more open-pored ascomata and smaller ascospores 

(see also under these species). Other similar, but 

chemically different Australian species include: 

T. rugatulum (nil), T. eungellaense and T. porinaceum 

(both norstictic acid). These taxa can be 

also easily distinguished by asci that do not have 

more than four spores. 

SPECIMENS EXAMINED – Australia, Queensland: Thornton 

Range, CREB rd. (to Cooktown): Stevens 19186 (GZU); 15 

km N of the Daintree River crossing, Hale 831017, 831622 (US); 5 km in from Daintree River crossing, Hale 

831523 (US). Mt. Windsor area, NW of Mossman: Km.45 on Mt. Windsor Rd., Hale 831303, 831432 (US); 5 

km W of new Forestry Camp, Hale 832066, 832210, 832246 (US). Davies Creek Rd. 17 km S of Kennedy 

Hwy., S of Davies Creek Falls NP., E of Mareeba, Hale 830998, 831054 (US). Mt. Tyson, 2 km WNW of Tully, 

Streimann 45596 (CANB). Dawson logging area, 24 km S of Koombooloomba turnoff, WSW of Tully, Hale 

830280 (US). 

Thelotrema triseptatum Mangold spec. nov. ined. 

Type: Australia, Queensland, Atherton Tablelands, Herberton Range, Hale 832261 (US-holotype). 

ETYMOLOGY – The epithet refers to the characteristic three-septate ascospores of this 

taxon. 



T. thesaurum. 

Thallus predominantly hypophloedal, very thin, epiphloedal parts up to 50 µm high, 

grayish. Surface dull to glittering, smooth to slightly roughened, continuous, unfissured to 

slightly fissured. Thallus covered by incontinuous protocortex up to 25 µm thick. Algal layer 

poorly developed, ±continuous, calcium oxalate crystals sparse, embedded in substrate, small 

to large. Vegetative propagules not seen. Ascomata inconspicuous, small, up to 300 µm in 

diam., roundish, apothecioid, solitary to marginally fused, distinctly immersed. Disc usually 

not visible from surface to rarely partly visible, pale flesh colored, epruinose. Pores small, up 

to c. 100 µm in diam., roundish to slightly irregular, upper parts of proper exciple visible from 

surface, free, entire to slightly split, off-white, incurved. Thalline rim margin moderately 

thick, entire to indistinctly split, roundish, often slightly sunken, concolorous with thallus, 

thalline rim not distinguishable from main thallus. Proper exciple free in the upper parts, thin, 

hyaline internally, yellowish-orange marginally, often with substrate layers incorporated, 

faintly amyloid (reddish). Hymenium up to c. 80 µm high, non-inspersed, distinctly 

conglutinated, paraphyses ±bent, interwoven, unbranched, tips thickened, lateral paraphyses 

present, inconspicuous, up to c. 10 µm long, columellar structures absent. Epihymenium 

indistinct, hyaline, without granules, with fine crystals. Asci 8-spored, tholus (moderately)


Fig. 176. Thelotrema triseptatum: growth habit (A), ascomata (B), ascospores (C) and ascospores 

showing amyloid reaction (D, E). A.-E.: US-holotype. Bar= A: 1.5 mm; B: 0.25 mm; C: 9 µm; D: 

5 µm; E: 4 µm. 

thick, moderately thin to moderatelt thick when 

mature. Ascospores uniform, small, transversely 

septate, cell walls thick, non-halonate, hyaline, 

moderately amyloid, fusi- to claviform, ends 

roundish to subacute, loci roundish to more often 

±angular, predominantly (roundish)cuboid, end 

cells conical, septae (moderately) thick, regular, 15- 

20 x 6-7 µm with 4 loci. Pycnidia not seen. 


C-, PD+ orange; containing stictic (major) and 

hypoconstictic (trace) acids. 

ECOLOGY AND DISTRIBUTION – The new species 

was collected on bark in a tropical rainforest at 

1100 m. It is only known from northern Queensland. 


T. triseptatum.


NOTES – Thelotrema triseptatum is characterized by a hypophloedal, ecorticate thallus, 

small, distinctly immersed ascomata, small, uniform three-septate, hyaline, moderately 

amyloid ascospores with thickened cell walls and septae, and the stictic acid chemosyndrome. 

Similar stictic acid containing Australian species include T. bicinctulum, T. alboolivaceum 

and T. porinoides. Thelotrema bicinctulum is distinguished by a true cortex and larger 

ascospores (up to 30 µm long with up to 10 loci). Thelotrema alboolivaceum has similar 

ascospores that are slightly larger (up to 23 µm long with up to 6 loci) and is further 

distinguished by a true cortex and larger, distinctly emergent, wide-pored apothecia with 

entirely free exciple. Thelotrema porinoides differs by having larger ascospores (up to 140 

µm, with up to 30 loci) with unthickened septae. 


Thelotrema sp. I 


Fig. 178. Thelotrema sp. I: growth habit (A) and ascoma (B). A., B.: Hale 831671. Bar= A: 1 mm; B: 

0.75 mm. 

NOTES – A single collection from Eungella National Park contains protocetraric acid and 

has small, roundish-elongate, regenerating ascomata that are ±concentrically arranged in a 

roundish, stroma-like structure that is limited by an erect to recurved 'thalline rim'. The 

ascomata have a layered (true/inner) thalline rim that is superficially slightly pale-reddish 

pigmented, an uncarbonized, fused proper exciple with lateral paraphyses up to 30 µm long, 

an up to c. 100 µm high hymenium with distinctly straight and parallel paraphyses with 

unthickened tips, and small, hyaline, transversely septate, non-amyloid ascospores up to c. 20 

x 6 µm in size with up to 6 loci (no mature ascospores seen). It is similar to Gyrotrema 

sinuosa (Sipman, 1992b; Frisch & Kalb, 2006) but differs in chemistry, the presence of lateral 

paraphyses, carbonization and smaller ascospores (hypoprotocetraric acid chemosyndrome, 

absent lateral paraphyses, carbonized proper exciple and ascospores up to 52 x 9 µm in size 

with up to 13 loci in G. sinuosa). Thelotrema sp. I lacks columellate structures and maybe be 

closer related to the T. schizoloma-group. 

SPECIMENS EXAMINED – Australia, Queensland, Eungella NP., NP. side rd. near Peases Lookout, off 

Darymple rd., Hale 831671 (US).


Thelotrema sp. II 


Fig. 179. Thelotrema sp. II: growth habit (A) and ascomata (B). A., B.: Lumbsch & Mangold 19128 c. 

Bar= A: 1 mm; B: 0.6 mm. 

NOTES – A single collection from Atherton Tableland could not be assigned to any known 

taxon, however, it is refrained from describing a new species since the specimen is too scanty. 

It is characterized by a moderately thick, smooth and slightly verrucose to rugose thallus with 

a large amount of small, scattered calcium oxalate crystals and covered by an incontinuous 

protocortex. It has medium sized apothecia with entire margins that become wide to gaping 

with age, fused to apically slightly detached proper exciple, clear hymenium up to 150 µm 

high with distinct lateral paraphyses and small (12-22 x 5-10 µm), submuriform, hyaline, nonamyloid 

to slightly amyloid ascospores with thick cell walls and predominantly ±irregular, 

angular, 4-6(7) x 1-2(3) loci. No secondary compounds could be detected by TLC. Chapsa 

kalbii from Africa and the Neotropics (type collection not seen) is similar, but differs by a 

more thinly thallus with rough surface, ascomata with split and rugged to lobed thalline rims, 

a lower hymenium (up to 70 µm high) and ascospores with less thickened walls and more 

roundish loci. 

SPECIMENS EXAMINED – Australia, Queensland, Atherton Tablelands, Curtain Fig Tree, Lumbsch & Mangold 

19128 c (F). 

Thelotrema sp. III 


NOTES – A single collection from Mossman Gorge (Northern Queensland) agrees well 

with the description for Myriotrema flavolucens from Venezuela, a species that is 

predominantly characterized by the presence of lichexanthone as the only secondary 

compound, immersed ascomata with layered margins and free proper exciple and small, 

hyaline, transversely septate, amyloid ascospores (Sipman, 1992). Unfortunately the type 

material of this taxon was not available for study. The Australian material is identical in the 

unusual chemistry and similar in its morphology and ascospores (up to 25 µm long). 

However, it differs by the presence of lateral paraphyses and an inspersed hymenium.


Fig. 180. Thelotrema sp. III: ascomata (A) and ascospore (B). A., B.: Mangold 35 k. Bar= A: 1 mm; 

B: 4 µm. 

SPECIMENS EXAMINED – Australia, Queensland, Daintree NP., Mossman Gorge Section, near Rex Creek 

Swing Bridge, Mangold 35 k (F). 

2. 9. 12. Topeliopsis Kantv. & Vezda Lichenologist 32: 347 (2000), emend. Kalb, Mycotaxon 

79: 320 (2001). Type species: Topeliopsis muscicola Kantv. & Vezda [= Topeliopsis 

muscigena (Stiz.) Kalb]. 

THALLUS – Crustose, muscicolous, corticolous, lignicolous humicolous or saxicolous, epito 

hyposubstratic, predominantly evanescent to ±thin, up to c. 50-200 µm high, rarely ±thick, 

up to c. 500 µm high, grayish to greenish or olive with yellow, brown or white tones. Surface 

dull to shiny or ceraceous, smooth to rarely rough or pruinose, continuous to rugose or 

verruculose, unfissured to rarely fissured or rimose. Prothallus absent or thin to indistinct and 

brownish. Thallus predominantly covered by an up to c. 10-30 µm thick, continuous to 

discontinuous protocortex, often becoming partly distinctly conglutinated forming a true 

cortex of irregular to periclinal hyphae to rarely without cortical structures or covered by a 

distinct, continuous, hyaline to yellowish true cortex consisting of periclinal to irregular 

hyphae up to c. 50 µm thick. Algal layer continuous to discontinuous, poorly to well 

developed, calcium oxalate crystals absent to abundant, small to large, scattered or clustered. 

Distinct medulla layer absent. Vegetative propagules are not known for the genus. 

ASCOMATA – Conspicuous to rarely inconspicuous, predominantly (moderately) large, up 

to c. 0.6-1.2 mm in diam., predominantly roundish to rarely slightly irregular, peri- to 

apothecioid, sessile to erumpent, solitary to slightly fused, non-regenerating, immersed to 

distinctly emergent, then hemispherical, urceolate or subglobose. Disc usually not visible 

from surface to partly visible, flesh-colored to brownish or grayish, rarely with red or orange 

tones, epruinose to rarely slightly pruinose. Pores predominantly tiny to wide, up to c. 100- 

600 µm in diam., predominantly ±irregular, ragged or distinctly star-shaped, rarely roundish, 

particularly in older ascomata, pore margin entire to ±split, proper exciple predominantly not 

visible from surface and pore margin formed by thalline rim, rarely becoming apically visible 

from surface, particularly in older or apically eroded ascomata, fused to rarely slightly 

detached, pale brown to reddish-brown to rarely whitish, grayish or pale yellowish, incurved. 

Thalline rim margin thin to thick, distinctly split to coarsely cracked, lobed, lacerate or 

eroded, thalline rim squamose to layered, often pruinose or exfoliating, whitish to off-white or 

concolorous with thallus, rarely with a reddish-brown base, incurved to erect. Proper exciple 

predominantly fused to apically exposed, moderately thick to very thick, rarely moderately 

thin, hyaline to pale yellowish internally, yellowish, orange, grayish or brownish marginally, 

rarely with substrate inclusions, apically sometimes more darkened and rarely covered by


grayish granules, predominantly ±distinctly amyloid towards the base and the subhymenium 

to rarely non-amyloid. Subhymenium thin to thick, concolorous with proper exciple. 

Hymenium non-amyloid, discoid to usually ±cupular, up to c. 90-250(300) µm high, noninspersed 

and clear in the Australian species (in T. meridensis inspersed), weakly to 

moderately, rarely strongly conglutinated. Paraphyses ±straight, parallel to slightly 

interwoven, unbranched, tips not thickened to slightly thickened. Lateral paraphyses present, 

conspicuous to rarely inconspicuous, not clearly separated from proper exciple, up to c. 15-40 

µm long, columellar structures absent. Epihymenium indistinct to moderately thick, hyaline to 

sometimes brownish, egranulose to rarely granulose. 

Asci 1-8-spored, non-amyloid, clavate, ascus walls not thickened, tholus thin to thick, not 

visible to thin at maturity. Ascospores uni- to quadriseriate, small to large, 12-210 x 4-55 µm, 

transversely septate to eumuriform. Cell walls thin to more often ±thick, smooth to sometimes 

±distinctly crenate to irregular, endospore in muriform ascospores thin to moderately thick, 

rarely non-halonate to ±distinctly halonate, sometimes only in younger stages, hyaline to 

rarely yellowish to brown, predominantly distinctly amyloid, rarely non-amyloid; oblong to 

ellipsoid or (bi-) fusiform to cylindrical or bacillar-fusiform, sometimes distinctly bent; with 

roundish to subacute, rarely distinctly acute ends; with 4-32 x 0-8 or multiple loci, loci 

roundish to ±angular, subglobose, oblong, lentiform, cuboid or ±irregular, with hemispherical 

to conical end cells, transverse septae thin to moderately thick, distinct to often indistinct or 

vanishing with age in densely muriform ascospores, regular to irregular, in some species 

ascospores generating ascoconidia in terminal stages (see below). 

PYCNIDIA – Not known for the genus, in T. elixii and T. muscigena old ascospores 

generating ascoconidia, oblong-irregular to bacilliform up to c. 4-7 x 1 µm. 

CHEMISTRY – Stictic acid chemosyndrome compounds present or absent. 

ECOLOGY AND DISTRIBUTION – The Australian Topeliospsis spp. were collected on 

epiphytic mosses, tree bark, dead wood and siliceous rock, rarely from peaty soil or plant 

debris, in altitudes ranging between 20 and 1500 m. The majority of species is found in 

rainforests and wet sclerophyll forests or in several forms of (sub)alpine woodlands and 

moorlands, in predominantly sub-tropical to cool-temperate, rarely tropical climates of Pacific 

northern Queensland to southern Victoria and Tasmania. As currently known, six of the 11 

species known in Australia are endemic (T. acutispora, T. darlingtonii, T. decorticans, T. 

elixii, T. kantvilasii, T. tasmanica), one is subantarctic (T. subdenticulata), three are 

paleotropical to paleotemperate (T. laceratula, T. muscigena, T. pseudoexanthismocapa) and 

one pantemperate (T. azorica). 

NOTES – This genus was introduced (Kantvilas & Vezda, 2000) with three species, 

characterized by sessile, exfoliating, perithecioid ascomata with a fused, pale to dark-brown 

or carbonized, cupular exciple, distinct lateral paraphyses and large, eumuriform, thin-walled, 

non-amyloid ascospores. Later, the genus was revised to its present circumscription (Kalb, 

2002). According to the uncarbonized type species T. muscigena, which has ascospores with 

±thickened walls (at least in younger stages), Topeliopsis was restricted to non-carbonized 

taxa with ascospores of the ‘thick-walled-type’ and several species were added (ibid., Frisch 

& Kalb, 2006a). Additionally, two species with a distinctly dark and carbonized proper 

exciple were excluded (Kalb, 2002) and are now placed in a separate genus (Melanotopelia 

rugosa, M. toensbergii). One of the newly added species (Frisch & Kalb, 2006a), T. 

darlingtonii, differs in having rather Thelotrema-like, erumpent ascomata with a less 

distinctly fused apical proper exciple and non-amyloid ascospores. However, in my 

observations, various species showed a heterogeneous habitus according to the nature of the


host substrate. For example in T. azorica, when muscicolous or saxicolous specimens have 

sessile, distinctly emergent, urceolate ascomata, whereas in specimen growing on soft wood 

the ascomata are erumpent, hemispherical and semi-emergent. Further, the ascospores in the 

examined collections of T. muscigena are often found ±distinctly pigmented. Consequently, I 

suggest to also accept species that differ from Topeliopsis s. str., characterized by sessile, 

±urceolate, fissured to eroded ascomata and hyaline to brown, amyloid ascospores, and also 

accept species with Thelotrema-like ascomata (as described above) and non-amyloid 

ascospores. Characters found in all taxa include a thick, rather pallid proper exciple with 

indistinctly separated, ±conspicuous lateral paraphyses and straight, parallel to only slightly 

interwoven paraphyses with never distinctly thickened tips. Similar are the members of the L. 

schizoloma-group and the genera Chapsa, Melanotopelia, Pseudoramonia and Thelotrema. 

The latter is distinguished by ascomata with different and ±distinctly free proper exciple, for 

differences to the other genera and the L. schizoloma-group see there. 

In the extended molecular phylogenic analysis (see also part 3) Topeliopsis s. str. forms a 

well supported monophyletic group, however, T. meridensis, which morphologically differs 

by an inspersed hymenium and a rather dark proper exciple, clusters with species of Chapsa 

and Thelotrema, the generic placement of this species needs further study. 


Topeliopsis acutispora Kalb 

Mycotaxon 79: 320 (2001). Type: Australia, Queensland, Cunninghams Gap NP., K. & A. Kalb 21901 

(CANB!-holotype, hb. Kalb!-isotype [as n. 21900]). 


Thallus predominantly muscicolous, often overgrowing adjacent debris and bark or 

exclusively corticolous, very thin to thin, c. up to 80µm thick, mainly episubstratic, in 

corticolous specimen partly hypophloedal, pale gray to grayish-green, appearing darker when 

growing on dark substrate. Surface dull to shiny, smooth, continuous to verruculose, 

unfissured. Thallus predominantly covered by an incontinuous protocortex up to c. 30µm 

thick, sometimes becoming distinctly conglutinated, forming a true cortex of irregular 

hyphae. Algal layer variable, mostly incontinuous and poorly developed, in corticolous 

specimen sometimes continuous and well developed, calcium oxalate crystals usually 

abundant and of variable size, scattered or in clusters. Vegetative propagules not seen. 

Ascomata conspicuous, (moderately) large, up to c. 1 mm in diam., roundish, peri- to 

apothecioid, sessile, predominantly solitary, in corticolous specimen moderately emergent 

and hemispherical, otherwise distinctly emergent and subglobose to urceolate. Disc usually 

not visible from surface, rarely becoming partly visible, pale flesh-colored, epruinose. Pores 

small to moderately wide, rarely gaping, up to c. 300 µm in diam., usually ragged and 

±irregular star-shaped, with distinctly split, incurved pore margin, proper exciple not visible 

from surface, in strongly eroded ascomata proper exciple margin becoming visible, then pore 

roundish to somewhat irregular with ±entire, incurved, pale brownish to reddish-brown 

margin. Thalline rim margin thin to moderately thick, with same color and structure as rest of 

thalline rim, thalline rim lacerate, coarsely pruinose to squamulose, often eroded, somewhat 

exfoliating with age and becoming slightly layered, conspicuously off-white to whitish. 

Proper exciple predominantly fused, sometimes tips becoming exposed, thick, hyaline to pale 

yellowish internally, (pale)orange to reddish-brown marginally, usually distinctly amyloid


towards the base and subhymenium. Hymenium up to c. 200 µm high, non-inspersed, 

moderately conglutinated, paraphyses unbranched, ±straight, parallel to slightly interwoven 

with unthickened to slightly thickened tips, lateral paraphyses present, not clearly separated 

from exciple, conspicuous, up to c. 30 µm long, columellar structures absent. Epihymenium 

indistinct, hyaline, without granules. Asci 8-spored, tholus moderately thick, thin when 

mature. Ascospores (moderately) large, transversely septate, rarely with a single longitudinal 

septum, cell walls thick to very thick, in younger ascospores distinctly halonate, hyaline, 

strongly amyloid, ±distinctly bent, bacillar-fusiform, with narrowed-rounded to (sub)acute 

ends, loci angular in younger stages, becoming roundish, subglobose to slightly lentiform to 

oblong with hemispherical to conical end cells, septae moderately thin, regular, 50-130(150) x 

10-15 µm with 19-32 loci. Pycnidia not seen. 


Fig. 181. Topeliopsis acutispora: growth habit (A), ascomata (B), ascospores (C, D) and ascospore 

detail (E). A.-D.: CANB-holotype; E.: Streimann 36551. Bar= A: 2.5 mm; B: 0.5 mm; C: 12.5 

µm; D: 40 µm; E: 8 µm. 

ECOLOGY AND DISTRIBUTION – Topeliopsis acutispora grows on epiphytic mosses, more 

rarely on tree bark or dead wood in warm temperate highland rainforests, cool temperate 

rainforests or wet sclerophyll forests in altitudes ranging from 50 to 1300m. It is common and 

wide-spread occurring from the Queensland/New South Wales border region to eastern 

Victoria and southern Tasmania and is endemic to Australia. 

NOTES – This taxon is characterized by a Topeliopsis-like habitus, large, transversely 

septate, thick-walled, hyaline, amyloid ascospores and the lack of secondary compounds. It is 

similar to T. subdenticulata but differs in ascospore morphology. Although smaller 

ascospores fall within the range of T. subdenticulata and a thorough examination of several


ascospores per specimen is needed for a reliably 

determination, the ascospores of T. subdenticulata 

never exceed 110 µm in length and do not have 

more than 25 loci. The ascospores in T. subdenticulata 

further appear to be more distinctly 

fusiform and less distinctly bent and the shape of 

loci tend to be stronger prolate, whereas the loci in 

T. acutispora ascospores are slightly angular. These 

differences are constant in all specimen examined, 

and no intermediate stages could be found. 

Topeliopsis patagonica from southern America is 

also similar, it is distinguished by slightly larger 

ascospores (up to 160 µm long) with more roundish 

ends, ascomata with smaller (up to 200 µm in 

diam.), distinctly dark pores and the presence of the 

stictic acid chemosydrome. Also similar is T. 

pseudoexanthismocarpa, for differences see under 

that species. 

SPECIMENS EXAMINED – Australia, Queensland: Carabeen 

Nature Refuge, 45 km E of Warwick, Lumbsch & Mangold 19175 d (F). Lamington NP., 9 km S of Beechmont, 

Tullawallah, Tibell 12741 (UPS). New South Wales: Border Ranges NP., Tweed Range, Brindle Creek Rd., 

along Tweed Range Scenic Drive, between The Pinnacle Lookout and Bar Mt., Wedin 3648 (UPS). New 

England NP., along fire trail from Park Entrance, Wedin 3606 (UPS). 35 km WSW Gloucester, Barrington Tops 

NP., Gloucester Tops, along Beech Forest Walk, Wedin 3498 (UPS). Mt. Wilson - Mt. Irvine Rd., 25 km NNE 

of Katoomba, Streimann 31601 (CANB). 5,4 km ESE of Katomba, Valley of Waters Creek, Tibell 12243 (UPS). 

Illawarra Hwy., Macquarie Pass, 6 km NE of Robertson, Thor 4788 (UPS). Monga NP., 27 km SE of 

Braidwood, Mangold 11 o, u, v (F). Mongarlowe River, Picnic area and river walk, Mangold 13 f (F). Misty 

Mountain Rd., 23 km NW of Batemans Bay: Currowan SF., Elix 22746 (CANB); Buckenbowra SF., Wedin 

3255 (UPS). Victoria: Drummer Creek, Drummer River Forest Reserve, 10 km E of Cann River settlement, 

Streimann & Pocs 65313 (B, CANB). Croajingolong NP., Double Creek Nature Walk, Mangold 9 c (F). 

Hensleigh Rd., 10,5 km SE of Bendoc, Elix 24106 (CANB). Result Creek, Bendoc-Orbost Rd., 13 km SW of 

Bendoc, Streimann 36551 (CANB). Errinundra NP.: 13 km SW Bendoc, along Gap Rd., at Result Creek, Wedin 

3361 (UPS); 30 km SSW Bendoc, Ellery Camp, Sassafras Creek side, Wedin 3386 (UPS). East Gippsland, 

Bonang rd., NE of Orbost, Mangold 8 k (F). Arte River, 30 km NE of Orbost: Elix 24247 (B); Glen Arte, Wedin 

3415 (UPS). Bemm River, Princes Hwy., 8 km SSW of Club Terrace, Elix & Streimann 19489 (B, CANB). 

Tasmania: Tahune picnic area along Huon river, 15 km W of Geeveston, 50 km WSW of Hobart, A. & M. 

Aptroot 23063 (ABL). 

Topeliopsis azorica (P. James & Purvis) Mangold comb. nov. ined. 

Bas.: Ramonia azorica P. James & Purvis, Arquipelago 11A: 11 (1993). Type: Azores, Faial, 14. Apr.1992, 

Purvis & P. James s.n. (AZ-holotype, BM!-isotype). 



T. acutispora. 

Thallus corticolous, saxicolous or muscicolous, variable due to different substrate 

structures, in saxicolous specimen very thin to incontinuous, predominantly endolithic, up to 

c. 50 µm high in ascomata area, otherwise very thin to thin, up to 100(150) µm high, epi- to 

hyposubstratic, grayish to greenish- or yellowish-gray. Surface dull to rarely slightly shiny, 

smooth to roughened, continuous to coarsely pruinose, unfissured to slightly fissured, often 

appearing distinctly fissured to areolate due to substrate structure. Thallus cover variable, 

predominantly covered by incontinuous protocortex up to 25 µm thick, rarely becoming 

distinctly conglutinated forming a true cortex of irregular to periclinal hyphae. Algal layer


Fig. 183. Topeliopsis azorica: growth habit (A, B), ascoma section (C), ascospore detail showing 

amyloid reaction (D), ascospore (E) and ascospore showing amyloid reaction (F). A., E.: BMisotpe; 

B., D., F.: Messuti & de la Rosa 4754; C.: Messuti 4485. Bar= A: 0.7 mm; B: 1 mm; C: 150 

µm; D: 12.5 µm; E, F: 25 µm. 

variable, predominantly ±incontinuous and poorly developed in some specimen patches of 

distinct and well developed algal layers areas present, calcium oxalate crystals not seen. 

Vegetative propagules not seen. Ascomata ±variable due to substrate structures, conspicuous 

to inconspicuous, moderately large, up to c. 800 µm in diam., roundish, perithecioid when 

young, becoming apothecioid with age, sessile to somewhat erumpent, solitary to marginally 

fused, distinctly emergent when growing on rock, mosses or hard bark, depressed-subglobular 

to (depressed-)urceolate, on soft bark only slightly emergent, hemispherical. Disc not visible 

from surface, rarely becoming partly visible, pale flesh-colored, epruinose to slightly 

pruinose. Pores small to moderately wide, up to c. 300 µm in diam., irregular to more rarely 

distinctly star-shaped, pore margin split, proper exciple not visible from surface. Thalline rim 

margin coarsely cracked to lacerate, thalline rim apically coarsely squamose to somewhat 

pruinose, whitish to off-white, in semi-emergent ascomata also concolorous with thallus, 

±exfoliating, incurved to slightly erect at pore area, erect to recurved and lobed towards the


margins. Proper exciple fused, thick, hyaline internally, pale yellow to pale yellowish-brown 

marginally, usually slightly to distinctly amyloid in internal exciple and subhymenium. 

Hymenium up to c.180 µm high, non-inspersed, moderately conglutinated, paraphyses 

straight, distinctly parallel, unbranched, with unthickened tips, lateral paraphyses present, 

conspicuous, not clearly separated from proper exciple, up to c. 30 µm long, columellar 

structures absent. Epihymenium indistinct and hyaline, sometimes with sparse, very small 

calcium oxalate crystals. Asci 1- to more rarely 2-spored, tholus moderately thin in young 

asci, not visible with maturity. Ascospores large, eumuriform, cell walls moderately 

thickened, endospore (moderately) thin, in younger stages with thin halo, hyaline, distinctly 

amyloid, in certain stages of maturity opaque amyloid, cell wall and endospore remaining 

non-amyloid, predominantly cylindrical to more rarely roundish-fusiform to very rarely 

roundish-bifusiform or reniform, with roundish to more rarely slightly narrowed-roundish 

ends, loci moderately large, roundish to somewhat angular, roundish-cuboid to irregular, 

transverse septae thin, distinct, regular, 70-160 x 15-45 µm with multiple loci. Pycnidia not 

seen. 

CHEMISTRY – Strain I: Thallus K+ yellowish to brown, C-, PD+ orange; containing stictic 

(major), α-acetylconstictic (major to trace), hypoconstictic, hypostictic (minors to traces), 

cryptostictic and hyposalazinic (traces) acids. Strain II: Thallus K-, C-, PD-; no secondary 



collected in Australia on siliceous rock in a warmtemperate 

rainforest at c. 700 m. (In Argentina it 

was collected on Nothofagus bark) It is rare in 

Australia known only from south-central New 

South Wales. This is the first report for Australia. It 

was described from the Azores and has since been 

recorded from Scotland 14 and Argentina (Lumbsch 

& al., 2008). 

N OTES – Topeliopsis azorica has a variable 

habitus. The Australian specimen has an almost 

evanescent thallus, in contrast, the corticolous 

collections from the Azores and Argentina have a 

distinct thallus. The ascomata in the specimens 

growing on rock as well as on bryophytes show a 

characteristic Topeliopsis-like morphology, with 

distinctly emergent, subglobose to urceolate, bright 

apothecia, whereas the specimen from Tierra del 

Fuego and parts of the Azores collections growing 


T. azorica. 

on soft bark show only slightly raised and comparably inconspicuous apothecia that are in 

some parts ±concolorous with the thallus. The specimens share several characters, such as 

large, hyaline, usually cylindrical ascospores with large loci and distinct transverse septae, a 

distinct to strong amyloid reaction, moderately thick, non-amyloid cell-walls, and a short 

hymenium. Hence they are included in a single species. Similar species include T. elixii, 

T. muscigena and T. laceratula. Topeliopsis muscigena can be distinguished by ascomata with 

reddish-brown bases, slightly larger, entirely amyloid ascospores that turn brownish with age 

14 The record from Scotland is not published in literature but is listed in the online archive for Scottish 

biodiversity (www.biodiversityscottland.gov.uk) by an unknown author.


and generate ascoconidia. For differences to T. elixii and T. laceratula see under these 

species. 

SPECIMENS EXAMINED – Strain I: Argentina: Neuquen, Messuti & de la Rosa 4754 (BCRU). 

Strain II: Australia, New South Wales, Valley of Waters, 5 km E of Katoomba (Blue Mnts. area), Thor 6114, 

6123 (S). Europe, Azores, Pico: S of Santa Luzia, 11. Jun.1978, James s.n. (BM); Lagoa do Caiado, 13. 

Jun.1978, James s.n. (BM). Argentina, Tierra del Fuego, Sierra Alvear, Messuti 4485 (BCRU). 

Topeliopsis darlingtonii Frisch & Kalb 

Lichenologist 38: (2006a). Type: Australia, Queensland, Darlington Range, Kalb 33979 (CANB!-holotype; 

hb. Kalb-, hb. Frisch-isotypes). 


Fig. 185. Topeliopsis darlingtonii: growth habit (A), ascospores (B, C) and ascospore detail (D). A.- 

D.: CANB-holotype. Bar= A: 1 mm; B: 25 µm; C: 12 µm; D: 7 µm. 

Thallus corticolous, (moderately) thin, up to 100 µm high, epi- to hypophloedal, pale 

yellowish-gray. Surface slightly shiny, smooth, continuous to slightly verruculose, unfissured. 

Thallus predominantly covered by a ±continuous protocortex up to c. 25 µm thick, in some 

parts becoming distinctly conglutinated forming a true cortex of predominantly irregular 

hyphae. Algal layer incontinuous and poorly developed, calcium oxalate crystals not seen. 


in diam., roundish, peri- to apothecioid, erumpent, solitary to marginally fused, immersed to 

moderately emergent, then irregularly depressed-hemispherical to irregularly depressedurceolate. 

Disc not visible from surface to rarely partly visible, grayish to pale grayish-


brown, epruinose to slightly pruinose. Pores tiny to (moderately) small, up to c. 100 µm in 

diam., predominantly irregular, pore margin entire to slightly split, incurved, proper exciple 

usually not visible from surface, in more open ascomata becoming partly visible, entire to 

slightly split, whitish, incurved. Thalline rim divided in two sections, inner parts coarsely 

cracked and (moderately) thick, lacerate and slightly layered, predominantly incurved, 

sometimes becoming slightly erect towards outer margins, off-white to whitish, outer parts 

resembling the 'true' thalline rim formed by main thallus, often indistinct and eroded or fused 

with inner thalline rim parts, coarsely cracked, incurved to erect, concolorous with thallus. 

Proper exciple predominantly fused, tips sometimes becoming ±free, thin to moderately thick, 

hyaline internally, pale yellowish to pale brownish marginally, apically covered by grayish 

granules, often with substrate particles incorporated, ±amyloid towards the base and 

subhymenium. Hymenium up to c. 150 µm high, non-inspersed, moderately conglutinated, 

paraphyses ±straight, parallel to slightly interwoven with unthickened to slightly thickened 

tips, lateral paraphyses present, not clearly separated from exciple, inconspicuous, up to c. 25 

µm long, columellar structures absent. Epihymenium indistinct to thin, hyaline, without or 

with few grayish granules. Asci 8-spored, tholus moderately thick, thin when mature. 

Ascospores moderately small to moderately large, transversely septate, cell walls 

(moderately) thin, often with thin halo, hyaline, non-amyloid to moderately amyloid (fide 

Frisch & Kalb, 2006a), oblong to fusiform, with narrowed-roundish to subacute ends, loci 

relatively large, roundish to slightly angular, lentiform to oblong, septae moderately thin, 

irregular, 35-60 x 6-10 µm with 10-16 loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+ yellow, C-, PD+ orange; containing stictic (major), constictic, 

α-acetylconstictic and consalazinic (minors) acids. 

ECOLOGY AND DISTRIBUTION – Topeliopsis 

darlingtonii grows on bark in a subtropical 

rainforest ('cool-temperate submontane rainforest' - 

fide Frisch & Kalb, 2006a) at 980 m. It is only 

known from southern Queensland. 

NOTES – It is characterized by only moderately 

emergent ascomata with coarsely split and 

moderately layered thalline margin, medium-sized, 

transversely septate, hyaline, thin-walled, nonamyloid 

(to weakly amyloid?) ascospores with 

relatively large and somewhat irregular loci and the 

stictic acid chemosyndrome. No similar species of 

Topeliopsis is known from Australia, the most 

similar species in Australia is the tropical 

Thelotrema capetribulense, which differs by a 

thicker, strongly verrucose/verruculose thallus 

surface, ascomata with distinctly free proper 

exciple, and thick-walled, clavate ascospores with 

smaller loci. The southern South American T. 


T. darlingtonii. 

patagonica is another stictic acid containing Topeliopsis with transverse septate ascospores, 

this taxon is readily distinguished by larger ascospores (up to 150 µm long). In contrast to the 

original species description (ibid.), I could not find an amyloid reaction of the ascospores and 

the exciple was found to be apically ±detached in several ascomata. 

SPECIMENS EXAMINED – Australia, Queensland, Darlington Range, K. & A. Kalb 33980 (hb. Kalb).


Topeliopsis decorticans (Müll. Arg.) Frisch & Kalb 

Lichenologist 38: 44 (2006). Bas.: Thelotrema decorticans Müll. Arg., Bull. Herb. Boissier 1: 54 (1893). 

Type: Australia, Victoria, Black Spur, 1888, Wilson '514' (G-lectotype, here selected; NSW!-isolectotype [see 

also notes]). 

Topeliopsis corticola Kalb, Mycotaxon 79: 322 (2001). Type: Australia, New South Wales, Blue Mnts. NP., 

Mnt. Wilson, K. & A. Kalb 20462 (CANB!-holotype). 


Fig. 187. Topeliopsis decorticans: growth habit (A), ascomata (B), ascoma section (C); ascospores 

(D) and ascospore showing amyloid reaction (E). A.-E.: Mangold 5 j. Bar= A: 1.5 mm; B: 1 mm; 

C: 175 µm; D, E: 12 µm. 

Thallus predominantly muscicolous, rarely lignicolous, corticolous or saxicolous, thin to 

more rarely moderately thin, up to c. 150 µm high, epi- to hyposubstratic, predominantly 

(pale)olive. Surface dull to slightly shiny, smooth, continuous to rarely somewhat 

verruculose, unfissured. Thallus cover variable, often incontinuous true cortex present, up to 

c. 30 µm thick, moderately dense, formed of highly conglutinated periclinal to irregular 

hyphae, sometimes uncovered or with incontinuous protocortex up to c. 20 µm thick. Algal 

layer continuous, well developed, calcium oxalate crystals lacking or very sparse, small, 

clustered. Vegetative propagules not seen. Ascomata conspicuous, moderately small to large, 

up to c. 1 mm in diam., roundish, perithecioid when young, becoming apothecioid with age, 

sessile, solitary to marginally slightly fused, predominantly distinctly emergent, subglobular 

when young, becoming urceolate to more rarely depressed-urceolate with age. Disc usually


not visible from surface, sometimes becoming partly visible, pale flesh-colored, epruinose. 

Pores small to moderately wide, rarely gaping, up to c. 500 µm in diam., irregular to starshaped 

with split pore margin in younger ascomata, in older ascomata often roundish with 

entire pore margin, proper exciple not visible from surface. Thalline rim margin coarsely 

cracked to slightly lacerate, in older ascomata often becoming ±entire and smooth, thalline 

rim apically coarsely squamose to somewhat pruinose, whitish to off-white, incurved. Proper 

exciple fused, thick, hyaline internally, pale yellowish to grayish marginally, internal exciple 

and subhymenial layers usually slightly to distinctly amyloid. Hymenium up to c. 200 µm 

high, non-inspersed, moderately conglutinated, paraphyses straight, distinctly parallel, 

unbranched, with unthickened tips, lateral paraphyses present, conspicuous, not clearly 

separated from proper exciple, up to c. 35 µm long, columellar structures absent. 

Epihymenium indistinct, hyaline, without granules or crystals. Asci 8-spored, tholus 

moderately thick in young asci, thin or not visible at maturity. Ascospores moderately small 

to moderately large, eumuriform, cell walls (moderately) thick, sometimes with crenate 

surface, endospore moderately thin to moderately thick, often with thin halo, hyaline, 

moderately to strongly amyloid, predominantly roundish-fusiform to reniform, rarely 

cylindrical, with roundish to narrowed-roundish ends, loci large, roundish to somewhat 

angular, subglobose to roundish-cuboid or slightly oblong, transverse septae thin to 

moderately thick, distinct, regular to slightly irregular, 30-70 x 10-25 µm with 8-16 x 1-6 loci. 



ECOLOGY AND DISTRIBUTION – The species 

grows on epiphytic mosses, more rarely on wood, 

tree bark or on siliceous rock in warm to cooltemperate 

rainforests and wet sclerophyll forests in 

altitudes ranging from 20 to 1500 m. It is common 

and wide-spread occurring from north-central New 

South Wales to southern Victoria and in Tasmania 

and is currently not known from outside Australia. 

N OTES – Topeliopsis decorticans is characterized 

by a Topeliopsis-like habitus, 8-spored asci 

with medium-sized, muriform, hyaline, thickwalled, 

amyloid ascospores and the absence of 

secondary compounds. Other Topeliopsis species 

with muriform ascospores have distinctly larger 

ascospores and 1-2-spored asci. 

The type in G was not available for study and 

thus the lectotype here selected is based on the 

duplicate in NSW dated from 1888. The collection 

mentioned in Müller’s original description is 


T. decorticans. 

specified with “ad Black Spur: Wilson, n. 514”, however, in NSW five different collections 

exist labeled ‘Wilson 514’ (dated 20., 21. and 23. March 1885, 1888 and January 1890). In 

recent publications (Galloway, 1985; Kantvilas & James, 1991; Frisch & Kalb 2006) no 

lectotype was selected, and the type in G is noted as holotype (in Galloway without date, in 

Kantvilas & James dated 1888, in Frisch & Kalb dated 1892). Since Kantvilas & James 

(1991) is the first publication that mentions the type collection with date, this specimen is 

selected as lectotype.


SPECIMENS EXAMINED – Australia, New South Wales: 35 km WSW Gloucester, Barrington Tops NP., 

Gloucester Tops, along Beech Forest Walk, Wedin 3483 (UPS). New England NP.: E of Armidale, Point 

Lookout, Weber & McVean L-49337 (CANB); Weeping Rocks Track, 34 km SW of Dorrigo, Streimann 65182 

(CANB). Monga SF., Forest River Rd., 5 km S of Monga, Elix 30243 (CANB). Monga NP., 27 km SE of 

Braidwood, Mangold 11 c, t (F). Victoria: Hensleigh Creek Rd., 18 km SSE of Bendoc, Elix & Streimann 21945 

(CANB). Hammonds Rd., Errinundra NP., 19 km SSW of Bendoc, Elix 24170 (CANB). Arte River, 30 km NE 

of Orbost, Elix 24241 (CANB). Bonang Rd., NE of Orbost, Mangold 8 l (F). Along Stony Creek, Colquhoun 

SF., 9 km E of Lakes Entrance, Elix 5364 (CANB). Snowfields, Baw Baw NP., Mt. Erica car park at start of 

walking track to Mt. Erica summit, Ford 2075896 (MEL). Korumburra, Wilson s.n. (NSW-603830). Wilson 

Promontory NP., Windy Saddle, Mangold 7 a (F). Bellell Creek, Marysville to Matlock rd., c. 3 km E of Lake 

Mountain turnoff, Filson 1048951 (MEL). Black Spur: 1885, Wilson '514' (NSW-603828); Jan. 1890, Wilson 

'514' (NSW-603829). Warburton, Dec. 1885, Wilson s.n. (NSW-603832). Sassafras, Wilson s.n. (NSW-603834). 

Ferntree Gully [W of Melbourne], Jan. 1892, Wilson s.n. (NSW-603833), 23. Dec.1899, Bastow s.n. (MEL- 

26207, -724548). Ottway Range, 14 km N of Apollo Bay, Mangold 5 i, j (F). Tasmania: Mt. Victoria Forest 

Reserve, along trail from Mt. Albert Rd., Wedin 3086 (UPS). Mt. Victoria Track, 08. Dec.1981, Kantvilas s.n. 

(BM-761893). Little Rapid River, 19. Feb.1982, Kantvilas s.n. (BM-761924). Anthony Rd.: Near turnoff to 

power station, Kantvilas 191/90 (HO); Near Lake Sandra Track, Kantvilas 40/89 (HO). 3 km S of Teepookana, 

Kantvilas 604/90 (HO). King William Saddle, 14 km SW of Derwent Bridge, Elix 26914 (CANB). Olga River at 

Line 7, Kantvilas 173/90 (GZU, HO). Styx River Valley, 27. Nov.1981, Kantvilas s.n. (BM-761886). 13 km W 

of Hobart, Mt. Wellington, Myrtle forest, Tibell 11002 (UPS). W of Tahune Bridge: In the Warra SST, Coupe 

WR 0081, Kantvilas 69/03 (HO); "Big Coupe", in the Warra SST, Kantvilas 218/98 (HO). Lyrebird Saddle 

around Mystery Creek, 5 km W of Lune River, 70 km SSW of Hobart, A. & M. Aptroot 23193 (ABL). 

Adamsons Rd. near Strathblane, 04. Dec.1981, Kantvilas s.n. (BM-761889). Bun Hill, Forestier Peninsula, 

Kantvilas 378/89 (HO). 

Topeliopsis elixii Frisch & Kalb 

Lichenologist 38: 40 (2006a). Type: Australia, Queensland, Darlington Range, Kalb 33979 (CANB!holotype; 

hb.Kalb-, hb.Frisch-isotypes). 


Thallus muscicolous to partly corticolous, thin, up to 100 µm high, predominantly 

episubstratic, pale yellowish-gray. Surface slightly shiny, smooth, continuous to slightly 

verruculose, non-fissured. True cortex present, up to c. 25 µm thick, continuous to 

incontinuous, consisting of periclinal hyphae. Algal layer continuous and well developed, 

calcium oxalate crystals not seen (fide Frisch & Kalb [2006a] present, forming a basal layer). 


in diam., roundish, apothecioid, erumpent, solitary, emergent, predominantly (depressed- 

)urceolate. Disc partly visible from surface, pale flesh-colored to reddish-brown, slightly 

pruinose. Pores becoming wide to rarely somewhat gaping, up to c. 500 µm in diam., irregular 

in younger ascomata, becoming ±roundish, at first opening as irregular cracks, in older 

ascomata with coarsely split to eroded pore margin, proper exciple not visible from surface. 

Thalline rim coarsely cracked to somewhat lobed, lacerate to eroded, often becoming slightly 

pruinose, concolorous with thallus or slightly brighter. Proper exciple fused, tips becoming 

somewhat exposed if thalline rim is strongly eroded, thick, not clearly separated from thalline 

margin, hyaline internally, grayish-brown marginally, ±amyloid towards the base and 

subhymenium. Hymenium up to c. 200 µm high, non-inspersed, moderately conglutinated, 

paraphyses ±straight, unbranched, parallel to slightly interwoven with unthickened to slightly 

thickened tips, lateral paraphyses present, not clearly separated from proper exciple, 

inconspicuous, up to c. 20 µm long, columellar structures absent. Epihymenium thin to 

moderately thick, hyaline, with grayish-brown granules. Asci 1-spored, tholus moderately 

thick to thin, not visible at maturity. Ascospores large, densely eumuriform, cell walls 

(moderately) thin, endospore thin, in younger stages with thin halo, hyaline, distinctly 

amyloid, oblong to ellipsoid with roundish to narrowed-roundish ends, loci roundish to


Fig. 189. Topeliopsis elixii: growth habit (A), ascoma (B), young ascospore (C), ascospore detail (D); 

ascoconidia producing ascospores (in E. showing amyloid reaction) (E, G), and conidia (F). A.-G.: 

CANB-holotype. Bar= A: 3 mm; B: 0.3 mm; C: 12.5 µm; D, G: 20 µm; E: 25 µm; F: 10 µm.


slightly angular, predominantly irregular, transverse septae only distinct in younger stages, 

thin and ±regular, becoming irregular, then vanish, finally developing ascoconidia (see 

below). 110-180 x 20-55 µm with multiple loci. Pycnidia not seen, ascoconidia developing in 

old ascospores, bacilliform, up to c. 7 x 1 µm. 

CHEMISTRY – Thallus K+yellow, C-, PD+orange; containing hypoconstictic, hyposalazinic 

(majors), hypostictic (minor) and α-acetylconstictic (trace) acids. 

ECOLOGY AND DISTRIBUTION – Topeliopsis elixii 

was collected on epiphytic mosses and adjacent tree 

bark in a subtropical rainforest ('cool-temperate 

submontane rainforest' - fide Frisch & Kalb, 

2006a) at 980 m. It is only known from southern 

Queensland. 

NOTES – This species is characterized by open, 

predominantly flattened-subglobose ascomata with 

visible, pruinose, flesh-colored discs and lacerateeroded 

margins, large, eumuriform, hyaline, 

amyloid ascospores that form ascoconidia, and the 

presence of hypoconstictic and hyposalazinic acids 

as major constituents. Ascoconidia also occur in the 

similar T. muscigena, which lacks secondary 

compounds and has larger (up to 210 µm), often 

slightly brownish ascospores. Other similar taxa 

include T. azorica, T. tasmanica (both known from 

Australia) and T. novae-zealandiae (known from 

Sri Lanka and New Zealand) which are all 

distinguished by a different chemistry. Topeliopsis azorica morphologically can be distinguished 

by slightly smaller ascospores (up to up to 160 µm) with larger loci and the lack of 

ascoconidia. Topeliopsis tasmanica and T. novae-zelandiae are readily distinguished by 

smaller (in both taxa up to 100 µm long), non- to faintly amyloid ascospores in 1-spored asci. 

SPECIMENS EXAMINED – Australia, Queensland, Darlington Range, K. & A. Kalb 33980 (hb. Kalb). 

Topeliopsis kantvilasii Mangold spec. nov. ined. 

Type: Australia, Tasmania, Mt. Field NP., Lake Dobson, Kantvilas & James 650/81 (BM-holotype, BMisotype). 

ETYMOLOGY – This species is dedicated to the Australian lichenologist Gintaras Kantvilas. 



T. elixii. 

Thallus lignicolous, thin to very thin, up to 100 µm high, hypo- to more rarely episubstratic, 

pale whitish- to yellowish-gray. Surface dull to slightly glittering, smooth, continuous, 

unfissured. Thallus covered by an incontinuous protocortex up to c. 20 µm thick. Algal layer 

incontinuous and poorly developed, calcium oxalate crystals not seen. Vegetative propagules 

not seen. Ascomata conspicuous, moderately large, up to c. 800 µm in diam., roundish, 

perithecioid when young, becoming apothecioid with age, erumpent, solitary to slightly 

marginally fused, immersed to somewhat emergent, then irregularly flattened-urceolate. Disc


Fig. 191. Topeliopsis kantvilasii: growth habit (A), ascomata (B), ascoma section (C) and ascospores 

(D, E). A.-E.: BM-holotype. Bar= A: 1.75 mm; B: 0.8 mm; C: 100 µm; D: 3 µm; E: 7 µm. 

usually not visible from surface, rarely becoming partly visible, dark-gray, pruinose. Pores 

predominantly tiny to small, rarely becoming moderately wide, up to c. 100(200) µm in 

diam., roundish to somewhat irregular, radiate split, pore margin area formed by apical proper 

exciple, pale yellowish to more rarely slightly grayish, incurved, proper exciple otherwise not 

visible from surface. Thalline rim thick, lacerate, often becoming eroded, sometimes slightly 

layered, concolorous with thallus to more often pale yellowish, ±exfoliating, incurved to erect 

internally, recurved in outer thalline rim layers. Proper exciple predominantly fused, tips 

becoming exposed due to erosion of thalline rim margin, (moderately) thick, hyaline to pale 

yellowish internally, yellowish to (pale) orange marginally, apically often brownish, slightly 

amyloid (reddish) in upper parts, more distinctly amyloid towards the base and subhymenium 

(purple). Hymenium up to c. 90 µm high, non- inspersed, moderately conglutinated,


paraphyses unbranched, ±straight, parallel to slightly interwoven with unthickened to slightly 

thickened tips, lateral paraphyses present, often not clearly separated from hymenium, 

conspicuous, up to c. 25 µm long, columellar structures absent. Epihymenium indistinct to 

thin, hyaline, in older ascomata with ±coarse grayish-brown granules, sometimes 

covered/intermixed with released ascospores. Asci 8-spored, tholus thin to absent. Ascospores 

small, transversely septate, in older stages loci sometimes becoming indistinctly 

longitudinally divided, cell walls (moderately) thick, non-halonate, soon becoming brownish 

to grayish-brown, non-amyloid to weakly amyloid, oblong to fusiform to somewhat 

claviform, with rounded to narrowed-roundish to more rarely subacute ends, loci large, 

becoming small and indistinct in older ascospores, roundish to angular, subglobose to oblong 

to irregular, septae moderately thin to moderately thick, ±irregular, 12-22 x 4-6 µm with 4-8 

(x2) loci. Pycnidia not seen. 

CHEMISTRY – Thallus K+yellow, C-, PD+orange; containing stictic (major), constictic 

(minor), α-acetylhypoconstictic and cryptostictic (traces) acids. 


kantvilasii grows on dead wood in cool-temperate 

subalpine rainforests and other forms of montane 

woodland in altitudes ranging from 1030 to 1230 

m. It is only known from western-central Tasmania. 

N OTES – This new species is readily 

distinguished from the currently known taxa in 

Topeliopsis by small, brown, transversely septate 

ascospores. It is further characterized by its 

yellowish, distinctly lacerate ascomata and the 

presence of the stictic acid chemosyndrome. A 

similar species is the tropical Chapsa platycarpa, 

which is distinguished by a corticate thallus and 

larger, Geaster-like ascomata (up to 2 mm in 

diam.) with distinctly free proper exciple. 

Topeliopsis kantvilasii is another atypical 

Topeliopsis species, resembling the taxa T. darlingtonii 

(with layered thalline rim and apically free 

proper exciple) or T. tasmanica (with non-amyloid 

ascospores), and is included in the genus only tentatively. 

SPECIMENS EXAMINED – Australia, Tasmania: Lake Dobson, Mt. Field NP., Elix 40010 (CANB). Walls of 

Jerusalem NP.: Lake Tyre, Kantvilas 124/87 (HO); 0.5 km NW of The Temple, Kantvilas 120/87 (HO). 

Topeliopsis laceratula (Müll. Arg.) Mangold comb. nov. ined. 

Bas.: Thelotrema laceratulum Müll. Arg., Flora 70: 399 (1887). Type: Australia, Trinity Bay (Cairns area), 

Sayer s.n. (G!-lectotype, here selected). 



T. kantvilasii.


Fig. 193. Topeliopsis laceratula: growth habit (A), ascomata (B), ascospore (C), ascospore detail (D) 

and same ascospore detail showing amyloid reaction (E). A., B.: G-lectotype; C.-E.: Lumbsch & 

Mangold 19132 k. Bar= A: 2 mm; B: 0.8 mm; C: 17.5 µm; D, E: 10 µm. 

Thallus epi- to hypophloedal, (moderately) thin, up to c. 200 µm high, dark to pale olive. 

Surface ceraceous, smooth, continuous to rugose, ±fissured. True cortex present, thick, up to 

50 µm thick, yellowish, continuous, consisting of predominantly irregular hyphae, sometimes 

with substrate inclusions. Algal layer well developed, continuous, calcium oxalate very 

abundant, small to large, often clustered, predominantly found in hypophloedal parts. 

Vegetative propagules not seen. Ascomata often conspicuous, moderately small to moderately 

large, up to c. 600(800) µm in diam., roundish to somewhat irregular, peri- to more rarely 

apothecioid, erumpent, solitary to sometimes fused, immersed to slightly emergent, then 

hemispherical. Disc not visible from surface. Pores tiny to (moderately) small, up to c. 200 

µm in diam., roundish to roundish-irregular or irregular, entire to more often split, proper 

exciple not visible from surface. Thalline rim margin thick to very thick, roundish to irregular, 

lacerate to coarsely squamose and layered, whitish to off-white to brownish due to 

protuberant substrate, incurved, outer thalline rim concolorous with thallus, often erect to 

slightly recurved. Proper exciple fused, moderately thin to moderately thick, hyaline to pale 

yellowish internally, brownish to grayish marginally, non-amyloid. Hymenium up to


c. 200 µm high, non-inspersed, weakly conglutinated, paraphyses parallel, unbranched, tips 

unthickened to slightly thickened, lateral paraphyses present, inconspicuous, short, up to c. 

15(25) µm long, not clearly separated from proper exciple, columellar structures absent. 

Epihymenium indistinct, hyaline, without granules or crystals. Asci 1(-2)-spored, tholus 

moderately thick, thin when mature. Ascospores (moderately) large, eumuriform, cell walls 

(moderately) thick, endospore moderately thin to moderately thick, covered by a thin, 

sometimes irregular halo, hyaline, loci strongly to opaque amyloid, cell walls and endospore 

non-amyloid, fusiform to bifusiform, with narrowed-roundish to more rarely subacute ends, 

loci usually large, roundish to slightly angular, subglobular to irregular, transverse septae 

(moderately) thin, distinct, regular, 70-160(170) x 20-35 µm with 20-30 x 1-8 loci. Pycnidia 

not seen. 



laceratula was collected in Australia on tree bark in 


1000 m. It is a common species occurring in 

northern to north-central Queensland. Besides 

Australia it is recorded from India (Patwardhan & 

Kulkarni, 1977) and Sri Lanka (newly reported 

here), having a paleotropical distribution. 


usually dark thallus with a waxy surface due to a 

thick true cortex that forms a distinct contrast to the 

bright, ±immersed ascomata with lacerate margins 

and small pores. Further it has large, eumuriform, 

hyaline ascospores with thick cell walls. The 

endospore remains hyaline in iodine, whereas the 

loci show a strong amyloid reaction in iodine. A 

similar species is T. azorica, it can be readily 

distinguished by a bright thallus that usually lacks a 

distinct, well developed cortex. Two similar 


T. laceratula. 

Thelotrema species are T. rugatulum and T. adjectum, for differences see under these species. 

Topeliopsis laceratula as well as T. pseudoexanthismocarpa (with transversely septate 

ascospores) are two tropical, corticolous members of Topeliopsis that have a ±thick and 

distinctly corticate thallus. However, both taxa agree well with the genus in all crucial 

characters. 


Portland Rds., Hale 832409 (US). Near Cedar Bay NP., rd. to Cooktown, Mangold 34 g (F). Thornton Range, 

CREB rd. (to Cooktown), about 5 km in from Daintree River crossing, NW of Mossman, Hale 832230 (US). 

Daintree NP., Mossman Gorge Section, near Rex Creek Swing Bridge, Mangold 35 zr (F). Near Lake Placid, S 

of Kuranda, K. & A. Kalb 21299 (hb. Kalb). Crystal Cascades, 5 km W of Cairns, Lumbsch & Mangold 19120 c, 

i (F). Atherton Tablelands: Lake Eacham NP., track around lake, Hale 831734 (US); Malanda Falls, Lumbsch & 

Mangold 19132 k (F); Zillie Falls, Falls Rd. NE of Millaa Millaa, Hale 831614, 831710 (US); Millaa Millaa 

falls, Lumbsch & Mangold 19139 s (F). About 5 km NW of Babinda at the bridge crossing of the Russell River, 

Hale 831104 (US). Babinda Boulders, NW of Babinda: Hale 831676 (US), Mangold 39 h, j, n, p, z (F). Francis 

Range, Woopen Creek Rd., 25 km in from Bruce Hwy., NW of Innisfail, Hale 832070, 832216, 832403 (US). 

4.5 km E of Cardstone on Tully River Rd. to Kareeya Power Station, W of Tully, Hale 830736, 831972 (US). 

Mt. Spec NP., Ridge on the Loop, on the Paluma Rd., WNW of Townsville, Hale 830705 (US). Eungella NP., 

Rosser Rd. entry point off Darymple rd., near Peases Lookout, Hale 831395, 832528 (US). Sri Lanka, 

Sabaragamuwa, Adams Peak, N of Gilimale, Hale 46391 (US).


Topeliopsis muscigena (Stiz.) Kalb 

Mycotaxon 79: 322 (2001). Bas.: Thelotrema muscigenum Stiz., Jahresber. St. Gall. nat. wiss. Ges. 1888/89: 

247 (1890). Type: South Africa, Cape Province, Aug. 1887, McOwan s.n. (Z-holotype; H-Nyl.-22438-, PREisotypes). 

Topeliopsis muscicola Kantv. & Vezda, Lichenologist 32: 348 (2000). Type: Australia, Tasmania, Quamby 

Bluff, Kantvilas 202/85 (HO!-holotype, hb. Vezda-isotype). 

Thelotrema indicum Hale, Mycotaxon 3: 177 (1975). Type: India, Tamil Nadu, Hale & Patwardhan 40185 

(US!-holotype, BSI-isotype). 


Fig. 195. Topeliopsis muscigena: growth habit (A), ascomata (B), ascoma section (C), ascoconidia 

producing ascospore (D), conidia (E) and ascospores (F, G). A., C., D.: HO-holotype of T. 

muscicola; B.: Streimann 42836; E.: Tibell 11447; F., G.: Mangold 5 d. Bar= A: 3 mm; B: 0.7 

mm; C: 125 µm; D: 10 µm; E: 5 µm; F: 25 µm; G: 35 µm. 

Thallus predominantly muscicolous, rarely lignicolous, corticolous or saxicolous (fide 

Kantvilas & Vezda, 2000), thin to more rarely moderately thin, up to c. 150 µm high, epi- to


hyposubstratic, pale yellowish gray to grayish green or (pale)olive. Surface dull to slightly 

shiny, smooth, continuous to somewhat verruculose, unfissured. Thallus cover variable, often 

true cortex present, continuous to incontinuous, up to c. 20 µm thick, consisting of periclinal 

hyphae, sometimes cortex structures absent or with incontinuous protocortex up to c. 10 µm 

thick. Algal layer variable, poorly to well developed, incontinuous to continuous, calcium 

oxalate crystals not seen. Vegetative propagules not seen. Ascomata conspicuous, moderately 

small to moderately large, up to c. 700 µm in diam., roundish, perithecioid when young, 

becoming apothecioid with age, sessile, solitary to marginally slightly fused, distinctly 

emergent, subglobular in younger stages, becoming urceolate with age. Disc not visible from 

surface to sometimes becoming partly visible, flesh-colored in younger ascomata, pale to dark 

brownish-gray in older ascomata, epruinose. Pores small to moderately wide, rarely gaping, 

up to c. 500 µm in diam., irregular to star-shaped, pore margin split, proper exciple not visible 

from surface. Thalline rim margin coarsely cracked to slightly lacerate, thalline rim apically 

coarsely squamose to somewhat pruinose, whitish to off-white, incurved, base smooth, 

±cylindrical, conspicuously reddish-brown. Proper exciple fused, thick, hyaline to pale 

yellowish internally, pale yellowish-brown marginally, apically sometimes dark-brown, 

internal exciple and subhymenium usually slightly to distinctly amyloid. Hymenium up to c. 

250(300) µm high, non-inspersed, moderately conglutinated, paraphyses straight, distinctly 

parallel, unbranched, with unthickened to slightly thickened tips, lateral paraphyses present, 

inconspicuous, not clearly separated from proper exciple, up to c. 30 µm long, columellar 

structures absent. Epihymenium indistinct and hyaline in younger stages, becoming 

(moderately) thick in older ascomata, brownish, without granules, rarely with some sparse, 

very small crystals. Asci 1- to rarely 2-spored, tholus moderately thick, becoming thin or not 

visible at maturity. Ascospores (very) large, densely eumuriform, cell walls in younger stages 

somewhat thickened, (moderately) thin at maturity, endospore thin, with (moderately) thin 

halo, hyaline in younger stages, becoming yellowish to pale brown at late maturity, mature 

ascospores distinctly to strongly or opaque amyloid, cylindrical to roundish-fusiform with 

roundish to slightly narrowed-roundish ends, loci ±small, predominantly roundish to slightly 

angular, subglobose to irregular-subglobose, transverse septae thin, distinct and ±regular in 

younger stages, becoming irregular and vanish with age, finally ascospores dissolving 

internally, generating ascoconidia (see below), 100-210 x 20-55 µm with multiple loci. 

Pycnidia not seen, old ascospores generating ascoconidia, oblong-irregular up to c. 4 x 1 µm. 




collected in Australia on epiphytic bryophytes, 

more rarely on wood or tree bark and on rocks (fide 

Kantvilas & Vezda, 2000) in various habitats 

ranging from cool-temperate rainforests to wet 

sclerophyll forests to several forms of (sub)alpine 

to coastal heath- or moorlands and shrubs in 

altitudes ranging from 40 to 1080 m. It is a 

moderately common species occurring in southern 

Victoria and Tasmania. It was also recorded from 

Africa, India and New Zealand (ibid.). 

NOTES – Topeliopsis muscigena is characterized 

by a typical Topeliopsis habitus with ascomata with 

smooth, cylindrical, reddish-brown base, single- to 


T. muscigena.


rarely double-spored asci with large, eumuriform, thin-walled, hyaline to brownish, amyloid 

ascospores that generate ascoconidia and the absence of secondary compounds. In 

thelotrematoid lichens, ascoconidia are thus far only known in Topeliopsis, another species 

with ascoconidia is T. elixii, for differences see under that species. Two other similar 

members of Topeliopsis with large, muriform, but always hyaline ascospores are known in 

Australia: T. tasmanica, which is readily distinguished by smaller (up to 100 µm), nonamyloid 

ascospores and the stictic acid chemosydrome and T. azorica (see under that species 

for differences). Also similar is Thelotrema tuberculiferum known from the Lesser Antilles 

and India. It differs by larger, perithecioid ascomata with thick, eroded-pruinose than split or 

squamose thalline rims with abundant calcium oxalate crystal inclusions and hyaline 

ascospores. As far as it could be judged from the examination of the poor type material in 

TUR-Vain., T. tuberculiferum is very close to Topeliopsis, however, no taxonomical 

consequences are proposed here until more specimens can be studied. 

SPECIMENS EXAMINED – Australia, Victoria: Ottway Ranges, Skene Creek-Colac rd., 13 km ENE of Apollo 

Bay, Streimann 42836 (B, CANB); 14 km N of Apollo Bay, Mangold 5 d, i (F). Tasmania: Ben Lomond NP., 32 

km E of Evandale, Tibell 11447 (UPS). New Zealand, South Island, Canterbury, Walker s.n. (Vezda, Lich. Rar. 

Exs. 459) (GZU). India, Tamil Nadu, Hale 50616 (US). 

Topeliopsis pseudoexanthismocarpa (Mont. & Bosch) Mangold comb. nov. ined. 

Bas.: Ocellularia pseudoexanthismocarpa Patw. & C. Kulk., Norw. J. Bot. 24: 130 (1977). Thelotrema 

pseudoexanthismocarpum (Patw. & C. Kulk.) Hale, Mycotaxon 11: 132 (1980). Type: India, Kerala, Anamalai 

Hills, Nagarkar & Gole 76308 (AMH-holotype, US!-isotype). 


Thallus predominantly epiphloedal to slightly hypophloedal, (moderately) thick, up to c. 

500 µm high, pale yellowish- to greenish-brown to pale olive. Surface waxy, smooth, 

continuous to verrucose, unfissured to somewhat rimose. True cortex present, up to c. 50 µm 

thick, yellowish, consisting of predominantly periclinal to more rarely irregular hyphae. Algal 

layer continuous but poorly developed, calcium oxalate crystals very abundant, of variable 

size, scattered or in clusters. Vegetative propagules not seen. Ascomata conspicuous, variable 

throughout development, (moderately) large, up to c. 0.8 mm in diam., roundish to somewhat 

irregular, peri- to apothecioid, erumpent, solitary to marginally fused, immersed to distinctly 

emergent, then subglobose to urceolate. Disc not visible from surface. Pores tiny to small to 

more rarely moderately wide, up to c. 200(400) µm in diam., roundish to slightly irregular, 

pore margin entire to slightly split, incurved, proper exciple not visible from surface, inner 

thalline rim layer (see below) might be confused with proper exciple. Thalline rim divided in 

two sections, inner parts coarsely cracked and relatively thin at first, becoming thick, 

distinctly lacerate, slightly layered, ±exfoliating, off-white or brighter then thallus, outer parts 

resembling the 'true' thalline rim formed by main thallus, only distinct in younger ascomata, 

in older ascomata becoming eroded and/or fused with inner thalline rim parts, coarsely 

cracked, erect to recurved, concolorous with thallus. Proper exciple fused to rarely apically 

somewhat exposed, thick to very thick, hyaline to pale yellowish internally, yellowish-brown 

to reddish-brown marginally, non-amyloid. Hymenium up to c. 200 µm high, non-inspersed, 

weakly conglutinated, paraphyses unbranched, slightly bent, parallel to slightly interwoven 

with unthickened to slightly thickened tips, lateral paraphyses present, not clearly separated 

from exciple, conspicuous, up to c. 20 µm long, columellar structures absent. Epihymenium 

indistinct, hyaline, without granules. Asci 4- to 8-spored, tholus thick, thin when mature. 

Ascospores (very) large, transversely septate, cell walls thick to very thick, usually with a 

±distinct crenate to slightly irregular surface, with thin halo predominantly in younger stages,


Fig. 197. Topeliopsis pseudoexanthismocarpa: growth habit (A), ascomata (B, C), ascospores (D, E) 

and ascospores showing amyloid reaction (F). A., B., D., E.: Hale 831148; C.: Hale 831536; F.: 

US-isotype. Bar= A: 1.25 mm; B: 0.5 mm; C: 0.6 mm; D, E: 20 µm; F: 23 µm. 

hyaline, strongly amyloid, ±bent, bacillar-fusiform, 

with narrowed-rounded to subacute ends, loci 

angular in younger stages, becoming roundish, 

subglobose to more often lentiform or oblong with 

hemispherical to conical end cells, septae moderately 

thin, regular to slightly irregular, 100-200 x 

10-20 µm with 12-30 loci. Pycnidia not seen. 




pseudoexanthismocarpa grows on tree bark in 

tropical rainforests. This rare species is restricted to 

northern Queensland. This is a new record for 

Australia. Previously it was known from India and 

Sri Lanka (Hale, 1981), hence being a paleotropical 

element. 


T. pseudoexanthismocarpa.


NOTES – This taxon is characterized by a thick, dark, waxy, distinctly corticate thallus, 

ascomata with bright, lacerate and two-parted margins, large, transversely septate, hyaline, 

thick-walled, amyloid ascospores and the absence of secondary products. Thelotrema kamatii, 

known from India and Sri Lanka, is probably a synonym, but the type material was not 

available for study. The two examined collections from Sri Lanka, identified and published 

under T. kamatii (Hale 50098, 50166; in Hale, 1981), belong to T. pseudoexanthismocarpa. A 

similar Australian species is T. acutispora, a temperate species that differs in a thin, ecorticate 

to indistinctly corticate thallus, sessile ascomata, and smaller ascospores (up to 130 µm long). 

Topeliopsis pseudoexanthismocarpa is morphologically somewhat deviant from most 

members of Topeliopsis, but similar to species as T. darlingtonii or T. laceratula (see also 

under this species). 

SPECIMENS EXAMINED – Australia, Queensland: Thornton Range, 5 km in from Daintree River crossing, NW 

of Mossman, Hale 831536 (US). About 5 km NW of Babinda at the bridge crossing of the Russell River, Hale 

831148 (US). Sri Lanka, Kalutara district, Hale 50098, 50166 (US). 

Topeliopsis subdenticulata (Zahlbr.) Frisch & Kalb 

Lichenologist 38: 44 (2006a). Bas.: Ocellularia subdenticulata Zahlbr., in Skottsberg (ed.), The Natural 

History of Juan Fernandez and the Easter Island 2, Botany: 329 (1924). Thelotrema subdenticulatum (Zahlbr.) 

Salisb., Lichenologist 5: 267 (1972). Type: Chile, Juan Fernandez Islands, Masafuera, C. & I. Skottsberg s.n. 

(W-holotype). 

Topeliopsis vezdae Kalb, Mycotaxon 79: 323 (2001). Type: Australia, Queensland, Styx River SF., Kalb & 

Williams 19199 (CANB!-holotype). 


Thallus predominantly muscicolous, often overgrowing adjacent debris and bark or 

exclusively corticolous, very thin to thin, c. up to 60(100) µm high, mainly episubstratic, in 

corticolous specimen partly endophloedal, pale gray to grayish-green, appearing darker when 

growing on dark substrate. Surface dull to shiny, smooth, continuous to verruculose, 

unfissured. Thallus predominantly covered by an incontinuous protocortex up to c. 30 µm 

thick, sometimes becoming distinctly conglutinated, forming a true cortex of irregular 

hyphae. Algal layer variable, mostly incontinuous and poorly developed, in corticolous 

specimen sometimes continuous and well developed, calcium oxalate crystals usually 

abundant and of variable size, scattered or in clusters. Vegetative propagules not seen. 

Ascomata conspicuous, (moderately) large, up to c. 1.2 mm in diam., roundish, peri- to 

apothecioid, sessile, predominantly solitary, in corticolous specimen moderately emergent 

and hemispherical, otherwise distinctly emergent and subglobose to urceolate. Disc not 

visible from surface to rarely becoming partly visible, pale flesh-colored, epruinose. Pores 

small to moderately wide, rarely gaping, up to c. 400(600) µm in diam., usually ragged and 

irregular to star-shaped, pore margin distinctly split, incurved, proper exciple not visible from 

surface, in strongly eroded ascomata apical proper exciple becoming visible, roundish to 

somewhat irregular, ±entire, incurved, pale brownish to reddish-brown. Thalline rim margin 

thin to moderately thick, with same color and structure as rest of thalline rim, thalline rim 

lacerate, coarsely pruinose to squamulose, often eroded, somewhat exfoliating with age and 

becoming slightly layered, conspicuously off-white to whitish. Proper exciple predominantly 

fused, in eroded ascomata tips sometimes apically exposed, thick, hyaline to pale yellowish 

internally, (pale)orange to reddish-brown marginally, usually distinctly amyloid towards the 

base and subhymenium. Hymenium up to c. 150 µm high, non-inspersed, moderately 

conglutinated, paraphyses unbranched, ±straight, parallel to slightly interwoven with 

unthickened to slightly thickened tips, lateral paraphyses present, not clearly separated from


exciple, conspicuous, up to c. 40 µm long, columellar structures absent. Epihymenium 

indistinct, hyaline, without granules or crystals. Asci 8-spored, tholus moderately thick, thin 

when mature. Ascospores (moderately) large, transversely septate, cell walls thick to very 

thick, often slightly crenate, in younger ascospores distinctly halonate, hyaline, strongly 

amyloid, ±bent, bacillar-fusiform to fusiform, with narrowed-rounded to subacute ends, loci 

angular in younger stages, becoming roundish, subglobose to lentiform with hemispherical to 

conical end cells, septae moderately thin, regular, 50-100(110) x 10-17 µm with 15-24(25) 


Fig. 199. Topeliopsis subdenticulata: growth habit (A), ascomata (B), ascospores (C) and ascospores 

showing amyloid reaction (D). A.: Elix & Streimann 19875; B.: Wilson s.n. (NSW-603831); C., 

D.: Lumbsch 10974 f. Bar= A: 2 mm; B: 0.4 mm; C, D: 20 µm. 


ECOLOGY AND DISTRIBUTION – Topeliopsis subdenticulata was collected in Australia predominantly 

on epiphytic mosses, more rarely on tree bark or dead wood in highland warm 

temperate rainforests, cool temperate rainforests or wet sclerophyll forests in altitudes ranging 

from 20 to 1150 m. It is a common and wide-spread temperate species occurring from 

northern New South Wales to southern Victoria and in Tasmania. Besides Australia it was 

reported from Chile and New Zealand (Salisbury, 1972) being a subantarctic element. 

NOTES – This taxon is characterized by a typical Topeliopsis-like habitus, (moderately) 

large, transversely septate, thick-walled, hyaline, amyloid ascospores and the absence of


secondary compounds. Kantvilas & James (1991) 

gave ascospore sizes up to 120 µm with 16-26 

septa for T. subdenticulata, probably due to the 

circumstance that some of the Tasmanian 

collections examined where in fact T. acutispora, a 

very similar species that virtually differs only by 

larger ascospores. For a detailed description of 

differences see under that species. Another similar 

species is T. darlingtonii, which is readily 

distinguished by smaller ascospores (up to 60 µm 

long, with up to 16 loci) and the presence of the 

stictic acid chemosyndrome. The South American 

T. patagonica was considered conspecific with T. 

subdenticulata (Frisch & Kalb, 2005 as 'O. 

patagonica Imshaug in ed.'), is here regarded as 

distinct based on larger ascospores (up to 150 µm 

long) and the presence of the stictic acid 

chemosyndrome. . 

SPECIMENS EXAMINED – Australia, New South Wales: Mt. 

Warning NP., track from summit to parking lot, Mangold 19 zd (F). C. 1 km W of Mt. Banda Banda, Kantvilas 

488/88 (NSW). Burraga Swamp, Allyn River Forest Park, Kantvilas 219/88 (NSW). Gloucester Tops, Kantvilas 

394/88 (NSW). Mt. Wilson - Mt. Irvine Rd., 25 km NNE of Katoomba, Streimann 31597 (CANB). Monga NP., 

27 km SE of Braidwood, Mangold 11 e (F). Rutherford Creek, 17 km SE of Nimmitabel, Elix & Kalb 40850 

(CANB). Victoria: East Gippsland: Result Creek, 13 km SW of Bendoc, Elix & Streimann 19875 (CANB); 

Errinundra River West Branch, 20 km S of Bendoc, Elix & Streimann 21966 (CANB); Errinundra NP., Elix 

24057 a (B), Elix 24058 (B, CANB), 24061(CANB), Elix & Streimann 19875 (CANB); Bonang rd., NE of 

Orbost, Mangold 8 g (F); Boggy Creek Gorge, near Nowa Nowa, Filson 16707 (MEL). Eastern Highlands, Baw 

Baw region, 30. Apr.1999, Ford s.n. (MEL-2075898). Tarra Valley, Ewers 6018 (CANB). Wilsons Promontory 

NP., on Sealers Cove hiking track, Mangold 6 p (F). Cement Creek Scenic Reserve, 6 km N of Warburton, 

Lumbsch 10974 f (F). Bellel Creek (near Maryville), 05. Oct.1983, Kantvilas s.n. (HO-113984). Black Spur, Jan. 

1890, Wilson s.n. (S-L328). Ottway NP.: 14 km N of Apollo Bay, Mangold 5 c, g (F); 10 km W of Apollo Bay, 

Mangold 2 c, e (F). Tasmania: Near Nelson Bay River, Kantvilas 629/84 (BM). Mt. Arthur, Wilson s.n. (NSW- 

603831). 6 km S of Warratah Hwy. jct. with the Murchison Hwy., Hale 68729 (US). Anthony Rd., Kantvilas 

36/89, 196/91 (HO). Algonkian Mtn., Kantvilas 134/90 (HO). Gordon Rd., c. 2 km N of Frodshams Pass, 

Kantvilas 102/97 (HO). W of Tahune Bridge, Warra SST., 19. May 1999, Kantvilas s.n. (HO-503376). Buckland 

Military Training Area, S of Bluestone Tier, Kantvilas 324/03 (HO). Ben Ridge Rd., Kantvilas 1116/81 (BM). 

Bun Hill, Forestier Peninsula, Kantvilas 339/89 (HO). 

Topeliopsis tasmanica (Kantv. & Vezda) Mangold comb. et stat. nov. ined. 

Bas.: Chroodiscus australis ssp. tasmanicus Kantv. & Vezda, Lichenologist 32: 334 (2000). Chroodiscus 

macrocarpus ssp. tasmanicus (Kantv. & Vezda) Galloway, Australas. Lichenol. 49: 17 (2001). Type: Australia, 

Tasmania, Mt. Geikie, Kantvilas 196/98 (HO!-holotype). 



T. subdenticulata. 

Thallus muscicolous to humicolous, very thin to entirely hyposubstratic, up to c. 80(100) 

µm high (in ascomata area), colorless to pale gray. Surface dull, pruinose, continuous, 

unfissured. Cortex structures absent in main thallus, basal parts of ascomata covered by 

incontinuous protocortex up to 20 µm thick. Algal layer incontinuous and poorly developed in 

main thallus, algal cells scattered in thin layer of somewhat gelatinous hyphae or within the 

substrate, continuous and well developed in basal parts of ascomata, calcium oxalate crystals 

not seen. Vegetative propagules not seen. Ascomata conspicuous, (moderately) large, up to


Fig. 201. Topeliopsis tasmanica: growth habit (A), ascomata (B), ascoma section (C), ascospore (D) 

and ascospore detail showing iodine accumulation (E). A.-E.: HO-holotype. Bar= A: 1 mm; B: 0.4 

mm; C: 130 µm; D: 17 µm; E: 6 µm. 

c. 1.2 mm in diam., roundish, perithecioid when young, becoming apothecioid, sessile, 

solitary to sometimes marginally fused, distinctly emergent, subglobose first, becoming 

(depressed-)urceolate. Disc partly visible from surface in older ascomata, orange-brown 

becoming dark grayish-brown with age, epruinose. Pores becoming wide with age, up to c. 

500 µm in diam., irregular to more often star-shaped to somewhat roundish, at first opening as 

±regularly radiating cracks, in older ascomata with deeply split and lobed pore margin, proper 

exciple not visible from surface. Thalline rim coarsely lacerate, distinctly lobed, often 

slightly layered, rarely somewhat eroded, formed of off-white to pale-brownish, slightly 

pruinose, coarse thallus bits, incurved to slightly erect with age. Proper exciple typical, fused, 

very thick, hyaline internally, pale yellowish to pale greenish-yellow marginally, weakly 

amyloid, pinkish, particularly towards the base and the subhymenium, subhymenium 

conspicuous, very thick, with same color as proper exciple. Hymenium up to c. 220 µm high, 

non-inspersed, strongly conglutinated, paraphyses straight, unbranched, parallel to slightly 

interwoven, tips unthickened to slightly thickened, lateral paraphyses present, not clearly 

separated from exciple, conspicuous, up to c. 40 µm long, columellar structures absent. 

Epihymenium moderately thick, orange-brown to brownish in older stages, without granules. 

Asci 2- to 4-spored, tholus thick, moderately thin when mature. Ascospores moderately large, 

densely eumuriform, cell walls and endospore thin, in younger stages with thin halo, hyaline, 

non-amyloid, ellipsoid to fusiform, with roundish to narrowed-roundish to more rarely


subacute ends, loci roundish to angular, irregular, transverse septae thin, indistinct and 

irregular in younger stages, vanishing with full maturity, 50-100 x 20-35 µm with multiple 


CHEMISTRY – Not tested, fide Kantvilas & Vezda (2000) containing stictic (major), 

constictic, cryptostictic (minors), 'norstictic' [=hypostictic?] and 'menegazziaic' [=α-acetylhypoconstictic?] 



tasmanica was collected in Australia on bryophytes, 

peaty soil and plant debris over several 

kinds of rock of alpine reaches in alpine heathlands 

in altitudes ranging from 800 to 1080 m. It is a rare 

species occurring in south-western Tasmania and is 

currently only known from there. 


evanescent to inconspicuous thallus, open, mainly 

subglobose to urceolate ascomata with ±visible, 

epruinose, ±brownish discs, regular star-shaped 

pores in younger stages that develop into lobed, 

incurved to somewhat erect margin in older 

ascomata. Further a thalline rim of coarse, bright, 

slightly pruinose thallus bits, a conspicuously thick 

proper exciple that reacts pink in iodine, 

moderately large, hyaline, non-amyloid, thinwalled, 

eumuriform ascospores in 2-4-spored asci 

and the presence of the stictic acid chemo- 

syndrome. Topeliopsis macrocarpa known from Argentina and New Zealand is a similar 

species, it was treated as a subspecies of T. tasmanica by Kantvilas & Vezda (2000) and 

Galloway (2001). In the original description of T. tasmanica (Kantvilas & Vezda, 2000) the 

number of spores per ascus and the ascospore sizes are noted as the only relevant differences 

to 'C. australis ssp. australis' (=T. macrocarpa). In contrast, I regard it as a distinct species 

based on differences in ascospore morphology, number of spores per ascus and additional 

morphological differences.T. macrocarpa has constantly single-spored asci with amyloid 

spores up to 150 µm long. Additionally the ascomata in T. macrocarpa differ by opening in 

irregular cracks, the thalline rim is more distinctly layered and consists of larger, more 

distinctly pruinose thallus pieces and the disc is never clearly exposed. 



T. tasmanica. 

2. 9. 13. Species of uncertain taxonomic placement: Leptotrema schizoloma-group 

NOTES – This species-group was first mentioned by Kantvilas & Vezda (2000) as 

“?Thelotremataceae species A” where it was listed as a single, highly variable taxon 

comparative to the newly introduced/revised genera with chroodiscoid, exfoliating apothecia 

(Chroodiscus, Pseudoramonia, Topeliopsis) from Tasmania. The study of the mentioned 

Tasmanian specimen and additional collections from the Australian mainland revealed, that 

the “species A” consists of two distinct taxa that where known and described as Leptotrema 

schizoloma and Thelotrema guadeloupense (see also under these species). Additionally, I


found three closely affiliated new species from Australia (T. parvizebrinum, T. subzebrinum, 

T. zebrinum) and a Neotropical species (T. refertum), differing in transversely septate and 

submuriform ascospores respectively, and a similar Neotropical species (T. cryptotrema) 

containing the psoromic acid chemosyndrome and with muriform ascospores up to 80 µm 

long, which is otherwise close to L. schizoloma. 

The group is characterized by erumpent, regenerating, peri- to apothecioid ascomata with a 

±distinctly layered margin, an apically free proper exciple that is marginally dark-brown to 

±distinctly carbonized and a usually concolorous subhymenium, distinct lateral paraphyses 

and a strongly conglutinated hymenium, distinctly straight and parallel paraphyses with 

predominantly not thickened tips. The asci and ascospores are heterogeneous. Several species 

have thickened ascus walls and lack a distinct tholus (otherwise a tholus is present), the 

ascospores are either hyaline or rarely slightly pigmented, non-amyloid or rarely distinctly 

amyloid, thin- or thick-walled. It is a conspicuously variable group with regards to the 

occurring secondary compounds, and a comparably high number of intraspecific chemotypes 

are found. Similar genera include Melanotopelia, Thelotrema and Topeliopsis, the two latter 

are readily distinguished by a more pallid, never distinctly carbonized exciple, Melanotopelia 

differs by sessile, never distinctly fissured or layered ascomata. 

I refrain from introducing a new genus for this group since further studies are needed to 

evaluate its systematic position. Preliminary molecular data could be gained for a single 

taxon, T. zebrinum (see part 3), which indicates a prominent, but poorly supported position in 

the molecular tree within the thelotremataceaen Graphidaceae. 


“Thelotrema” guadeloupensis Hale 

Phytologia 26: 416 (1973). Type: Guadeloupe, Parc National de Guadeloupe, Hale 31633 (US! -holotype). 


Thallus variable, thin to very thin, epi- to hypophloedal or partly entirely hypophloedal, up 

to c. 150 µm high, rarely pale yellowish-gray to more often pale grayish-green to (pale) olive. 

Surface dull to shiny, smooth, continuous to verrucose, often distinctly fissured. Thallus cover 

variable, usually covered by a continuous to incontinuous protocortex up to c. 20 µm thick, 

sometimes becoming distinctly conglutinated, forming a true cortex of irregular to periclinal 

hyphae. Algal layer continuous and well developed, calcium oxalate crystals sparse, small to 

large, predominantly clustered. Vegetative propagules not seen. Ascomata variable, usually 

inconspicuous, moderately large, up to c. 800 µm in diam., roundish, perithecioid to 

apothecioid at maturity, erumpent, solitary to more fused, regenerating, predominantly 

±emergent, (irregular-)hemispherical to (irregular-)urceolate. Disc usually not visible from 

surface, rarely becoming partly visible, pale grayish, epruinose. Pores small to moderately 

wide to rarely wide, up to c. 300(400) µm in diam., roundish to irregularly angular, formed by 

apical margin of proper exciple, pore margin/visible part of proper exciple ±split, fused to 

indistinctly free, off-white to pale brownish, incurved. Thalline rim (moderately) thick, 

becoming distinctly layered with proceeding succession of ascomata generations, coarsely 

split to lacerate, more rarely somewhat eroded, (dark) brownish to blackish, ±distinctly 

contrasted by off-white layers, outer thalline rim concolorous with thallus, in distinctly 

layered thalline rims exfoliating, incurved to erect in inner layer(s), to erect to slightly 

recurved in outer layer(s), in un-layered thalline rims predominantly incurved. Proper exciple


Fig. 203. Thelotrema guadeloupensis: growth habit (A, C), ascomata (B), ascoma section (D) and 

ascospores (E, F). A., E.: US-holotype; B.: Kantvilas 572/92; C.: Kantvilas 552/88; D.: Wedin 

3273; F.: A. & K. Kalb 20461. Bar= A: 1.25 mm; B: 0.35 mm; C: 0.75 mm; D: 200 µm; E: 6.5 µm; 

F: 15 µm. 

predominantly apically free, moderately thick, hyaline internally to (dark-)brown to 

carbonized marginally, usually with substrate inclusions, distinctly amyloid at the base. 

Hymenium up to c. 200 µm high, non-inspersed, strongly conglutinated, paraphyses straight 

to slightly bent, ±distinctly parallel, unbranched, tips unthickened, lateral paraphyses present, 

conspicuous, up to c. 30 µm long, true columella absent, columella-like structures sometimes


present in newly developing ascomata generations (=raised subhymenial layer) or in fused 

ascomata, upper subhymenial layer dark-brown to carbonized, sometimes followed by a 

hyaline layer of newly developing hymenium. Epihymenium indistinct, hyaline, without 

granules or crystals. Asci 4 to 8-spored, tholus and lateral ascus walls (moderately) thick, thin 

when mature. Ascospores moderately small to moderately large, eumuriform, cell walls and 

endospore thin, non-halonate, hyaline, non-amyloid, predominantly ellipsoid to fusiform with 

narrowed-roundish to sub-acute ends, loci roundish to angular, predominantly roundishcuboid 

to cuboid or irregular, transverse septae thin, distinct, regular, (20)30-60(70) x 10-25 

µm with 8-20 x 1-8 loci. Pycnidia not seen. 

CHEMISTRY – Strain I: Thallus K+yellow, C-, PD+orange; containing stictic (major), 

constictic (major to minor), and α-acetylhypoconstictic (trace) acids. Strain II: Thallus K-, C-, 

PD-; containing the 'cinchonarum unknown' compound (major), [dark-gray spot after 

charring, Rf 2/7/3 (solv. syst. A/B'/C)]. 


guadeloupensis was collected in Australia on tree 

bark in warm- to cool-temperate, rarely sub-tropical 

highland rainforests in altitudes ranging from 500 

to 1000 m. It is rare but wide-spread occurring in 

northern to southern New South Wales and in 

(south-)western Tasmania. This is the first report of 

the species from Australia. It was previously 

recorded from Tasmania as “spec. A” (Kantvilas & 

Vezda, 2000). So far it was only known from 

Guadeloupe. 

NOTES – This taxon is readily distinguished by 

the regenerating, distinctly layered and carbonized 

ascomata (at least in older stages), medium-sized, 

muriform, thin-walled, hyaline, non-amyloid 

ascospores and the presence of either the stictic 

acid chemosyndrome or 'cinchonarum unknown'. It 

was treated and published under the provisional 

name 'Spec. A' (Kantvilas & Vezda, 2000), which 

also included L. schizoloma, a similar species that differs in larger ascospores (up to 130 µm 

long) in 1-4-spored asci. The only other known similar species is T. zebrinum that differs in 

having transversely septate ascospores. 

SPECIMENS EXAMINED – Strain I: Australia: New South Wales: Dorrigo NP., Sassafras Creek Track, 

Mangold 25 l (F). Blue Mountains NP.: Mt.Wilson, K. & A. Kalb 20461 (hb. Kalb); Near Blackheath, Hale 

58595 (US). Monga NP.: 18 km SE of Braidwood, Wedin 3273 (UPS); 27 km SE of Braidwood, Mangold 11 c, 

h, i, m, n, p, q, r (F). Tasmania: Mt.Murchison, Anthony Rd., Kantvilas 572/92 (HOB). Mt.Dundas Track, 

Kantvilas 552/88 (B, HOB). Lake St.Clair NP., Cradle Mnt., Pelion Plains, Kantvilas 217/92 (HOB). 

Strain II: Australia, Tasmania: Mt.Murchison, Anthony Rd., Kantvilas 239/93 (HOB). Walls of Jerusalem 

NP., Little Fisher River, Kantvilas & James 463/84 (HOB). Mueller Rd., 3 km W of Styx Rd., Kantvilas 646/84 

(HOB). 

“Thelotrema” parvizebrinum Mangold spec. nov. ined. 


T. guadeloupensis. 

Type: Australia, Queensland, Mt. Chalmynia logging area, c. 15 km from Bruce Hwy, W of Innisfail, Hale 

832635 (US-holotype).


ETYMOLOGY – The epithet refers to the similar species T. zebrinum and the smaller 

ascomata and ascospores (from lat. parvus =small). 


Fig. 205. Thelotrema parvizebrinum: growth habit (A), ascoma section (B) and ascospore showing 

amyloid reaction (C). A.-C.: US-holotype. Bar= A: 0.2 mm; B: 75 µm; C: 5 µm. 

Thallus thin to moderately thick, epi- to hypophloedal, up to c. 200 µm high, rarely pale 

yellowish-gray to more often grayish-green to pale olive. Surface dull, smooth to slightly 

roughened, distinctly to strongly verruculose, fissured. Thallus covered by an incontinuous to 

more rarely continuous protocortex up to c. 20 µm thick. Algal layer continuous to somewhat 

incontinuous, well to moderately well developed, calcium oxalate crystals variable, 

sometimes sparse, usually frequent, in clusters, often also embedded in substrate tissue. 

Vegetative propagules not seen. Ascomata inconspicuous, small, up to c. 200 µm in diam., 

roundish, becoming apothecioid, erumpent, solitary, immersed to rarely slightly emergent, 

then depressed-hemispherical. Disc usually not visible from surface, very rarely becoming 

partly visible, whitish, epruinose. Pores tiny to small, up to c. 60(100) µm in diam., 

predominantly irregular, formed by apical margin of proper exciple, pore margin/visible part 

of proper exciple ±split, fused to indistinctly free, off-white to pale brownish, incurved. 

Thalline rim moderately thick to moderately thin, becoming slightly layered in older stages, 

split to slightly lacerate, sometimes eroded, (dark) brownish, sometimes indistinctly 

contrasted by off-white to pale brownish layers, outer thalline rim concolorous with thallus, 

incurved to erect. Proper exciple becoming apically to almost entirely free, moderately thick, 

hyaline to pale yellowish internally, grayish to (dark) brown or slightly carbonized 

marginally, sometimes with substrate inclusions, often amyloid in lower parts. Hymenium up 

to c. 100 µm high, non-inspersed, strongly conglutinated, paraphyses straight, parallel to 

slightly interwoven, unbranched, with unthickened to slightly thickened tips, lateral 

paraphyses present, usually very inconspicuous, up to c. 15 µm long, columellar structures 

absent. Epihymenium indistinct to thin, hyaline, rarely with fine grayish to colorless granules. 

Asci 8-spored, tholus (moderately) thick, thin or not visible at maturity. Ascospores small,


transversely septate, cell walls (moderately) thick, in younger stages with thin halo, hyaline, 

distinctly to strongly amyloid, oblong to fusiform or somewhat claviform, with roundish to 

subacute ends, loci roundish, subglobose to slightly lentiform, septae moderately thin to thick, 

regular, 12-20 x 4-6 µm with 4-7(8) loci. Pycnidia not seen. 



parvizebrinum was collected in Australia on tree 

bark in tropical rainforests in altitudes ranging from 

100 to 1200 m. It is rare and currently only known 

from northern Queensland. 

NOTES – The new species is characterized by a 

verruculose thallus, small, usually immersed 

ascomata with often slightly layered, brownish to 

slightly carbonized proper exciple, small, transversely 

septate, hyaline, amyloid ascospores with 

thickened cell walls and the absence of secondary 

compounds. It can be distinguished from T. 

zebrinum by the smaller ascomata with less 

distinctly layered margins, the smaller ascospores 

(up to 80 µm long with up to 22 loci in T. 

zebrinum) and the absence of secondary 

compounds (stictic and/or protocetraric acid 

chemosyndrome in T. zebrinum). Thelotrema 

defossum is another similar species, which differs in 

having larger ascospores (usually up to 30 µm long with up to 11 loci) with less thickened 

cell walls, a distinct halo, more angular, flattened loci. Thelotrema refertum known from 

Panama and Brazil 15 is also similar to T. parvizebrinum, but differs by a continuous to rimose, 

corticate thallus and the hypoprotocetraric acid chemosydrome. 

SPECIMENS EXAMINED – Australia, Queensland: Thornton Range, 5 km in from Daintree River crossing, NW 

of Mossman, Hale 831714 (US). Mt. Windsor logging area, 9 km from rd. to old Forestry Camp and the main 

rd., NW of Mossman, Hale 832424 (US). Mt. Lewis Rd. 4 km N of Kennedy Hwy., W of Mossman, Hale 

831235, 832549 (US). 

“Leptotrema” schizoloma Müll.Arg. 

Nuov. Giorn. Botan. Ital. 21: 49 (1889). Type: Argentina, Tierra del Fuego ['Fuegia'], Ushuaia, Beagle 

Channel, 1885, Spegazzini 101 (ex Herb. Jatta) (G!-lectotype, here selected). 

Thelotrema hypomelaenum Müll. Arg., Bull. Herb. Boissier 3: 314 (1895). Type: Australia, New South 

Wales, Knight 16 pr.p. (G!-lectotype, here selected). 



T. parvizebrinum. 

Thallus variable, epi- to hypophloedal, very thin to moderately thick, up to c. 300 µm high, 

pale gray to pale yellowish-gray or pale grayish-green to yellowish-olive. Surface dull to 

slightly shiny, smooth to rough, continuous to strongly verrucose or verruculose, often 

distinctly fissured. Thallus predominantly covered by a ±continuous protocortex, up to 

15 Unpublished collection in US, newly reported here: Brazil, Amazonas, Rio Uatuma, Buck 2943 (US).


Fig. 207. Leptotrema schizoloma: growth habit (A, B), ascomata (C, D), ascoma margin (E), 

ascospores (F, G) and ascospore showing amyloid reaction (H). A., F., G.: A. & K. Kalb 20560; 

B.: Hale 832271; C.: Spegazzini s.n.; D.: Lumbsch 8678 c; E.: G-lectotype; H.: Mangold 22 r. 

Bar= A: 1 mm; B: 0.8 mm; C: 0.5 mm; D: 0.35 mm; E: 110 µm; F: 16 µm; G: 15 µm; H: 24 µm. 

c. 20 µm thick, rarely becoming distinctly conglutinated forming a true cortex of periclinal 

hyphae. Algal layer continuous and usually well developed, calcium oxalate crystals 

abundant, small to large, solitary or in clusters. Vegetative propagules not seen, isidia-like 

structures sometimes present in strongly verruculose specimen. Ascomata variable, 

conspicuous, large, up to c. 1.2 mm in diam., roundish, perithecioid in younger stages,


becoming indistinctly apothecioid with age, erumpent, regenerating, solitary to more rarely 

fused, predominantly ±emergent, (irregular-)hemispherical to (irregular-)depressed-urceolate. 

Disc not visible from surface to very rarely becoming partly visible, pale grayish to whitish, 

epruinose. Pores tiny to (moderately) small, up to c. 250 µm in diam., roundish to irregular, 

formed by apical margin of proper exciple, pore margin/visible part of proper exciple ±split, 

fused to indistinctly free, off-white to pale brownish, incurved. Thalline rim (moderately) 

thick, becoming distinctly layered with proceeding succession of ascomata generations, 

coarsely split to predominantly lacerate, more rarely somewhat eroded, (dark) brownish to 

blackish, ±distinctly contrasted by off-white layers, outer thalline rim concolorous with 

thallus, in distinctly layered thalline rims exfoliating, incurved to erect in inner layer(s), to 

erect to slightly recurved in outer layer(s), in un-layered thalline rims predominantly 

incurved. Proper exciple moderately thick, predominantly apically free, predominantly 

entirely (dark-)brown to usually distinctly carbonized, sometimes with substrate inclusions, 

distinctly amyloid at the base. Inner subhymenial layer dark-brown to carbonized, sometimes 

followed by a hyaline layer of newly developing hymenium. Hymenium up to c. 200 µm 

high, non-inspersed, strongly conglutinated, paraphyses straight to slightly bent, ±distinctly 

parallel, unbranched, tips unthickened, lateral paraphyses present, conspicuous, up to c. 30 

µm long, true columella absent, columella-like structures sometimes present in newly 

developing ascomata generations (=raised subhymenial layer) or in fused ascomata. 

Epihymenium indistinct, hyaline, without granules or crystals. Asci 1-4(6)-spored, tholus and 

lateral ascus walls (moderately) thick, thin when mature. Ascospores (moderately) large, 

densely eumuriform, cell walls and endospore slightly thickened in younger stages, becoming 

thin at maturity, non-halonate, hyaline to yellowish or rarely pale brownish in late maturity or 

when deceased, non-amyloid to faintly amyloid in older ascospores, predominantly oblong to 

ellipsoid with roundish to narrowed-roundish ends, loci predominantly ±angular, 

predominantly irregular, becoming small with age, transverse septae thin, distinct in younger 

stages, becoming indistinct with age, predominantly ±irregular, (40)60-120(130) x 20-40 µm, 

with multiple loci. Pycnidia not seen. 

CHEMISTRY – Strain I: Thallus K-, C-, PD-; no secondary compounds detectable by TLC. 

Strain II: Thallus K+ yellow becoming deep red, C-, PD+ orange; containing salazinic acid 

(major). 

ECOLOGY AND DISTRIBUTION – Leptotrema 

schizoloma occurs on bark and wood in cool- to 

warm-temperate rainforests, more rarely in 

(sub)tropical highland rainforests, in altitudes 

ranging from 100 to 1500 m. It is a moderately 

common and wide-spread species occurring along 

the east coast from high altitudes in northern 

Queensland to New South Wales and also 

Tasmania. It is also known from Argentina and 

New Zealand (Lumbsch & al., 2008), being an 

subantarctic element that extends to the subtropics 

at high elevations. 

NOTES – This taxon is readily distinguished by 

the regenerating, distinctly layered and carbonized 

ascomata (at least in older stages), large, densely 

eumuriform, thin-walled, hyaline to slightly 

pigmented, non- to faintly amyloid ascospores and 


L. schizoloma.


either the absence of secondary compounds (strain I) or the presence of salazinic acid (strain 

II). Similar is T. guadeloupensis, for differences see under that species. 

SPECIMENS EXAMINED – Strain I: Australia: Queensland: Bellenden Ker NP., below the cable car line near 

the summit, Stevens 22033 (GZU). Lamington NP., Main Border Track out of O'Reillys, Hale 832271 (US). 

New South Wales: Border Ranges NP., NE of Wiangaree, Hafellner 16592 (GZU). Mt. Warning NP., track from 

summit to parking lot, Mangold 19 c, m, u, zg, zl, ze, zn (F). Dorrigo NP.: Never Never Picnic Area and 

Rosewood Creek Track, Mangold 24 c (F); Sassafras Creek Track, Mangold 25 o, s (F). Nightcap Forest Drive: 

Gibbergunyah Roadside Reserve, Whian Whian SF., W of Mullumbimby, Hale 831692 (US); 1 km W of 

Minyon Falls, N of Lismore, Hale 832193 (US). Nightcap NP., Mt. Nardi/Mt. Matheson Track, Mangold 22 r 

(F). Barrington Tops, Stewarts Brook SF., Lumbsch & Filson 8678 c (CBG, F). Doyles River SF. on Oxley 

Hwy., 95 km SE Walcha, Hale 58638, 58566 (US). Mill Creek, 50 km NW of Sydney, K. & A. Kalb 34268, 

34270 (hb. Kalb). Royal NP., Bola Creek, K. & A. Kalb 20560, 21694 (hb. Kalb). Blue Mts., Below Bridal Veil 

Falls, nr. Blackheath, Hale 58539, 58619, 58579, 58547 (US). Tasmania: Mt. Murchison, Anthony Rd., 

Kantvilas 202/93 (HOB). Argentina, Isla de los Estados ['Staten Island '], Port Cook, 1888, Spegazzini s.n. (G). 

Strain II: Australia, Tasmania: Walls of Jerusalem NP., un-named lake, 1.5 km W of Chalice Lake, Kantvilas 

95/99 (HOB). Hartz Mtns. NP., Lake Osborne Track, Kantvilas & James 496/81 (HOB). 

“Thelotrema” subzebrinum Mangold spec. nov. ined. 

Type: Australia, New South Wales, Mt. Warning NP., track from summit to parking lot, Mangold 19 e 

(CANB-holotype; NSW-, F-isotypes). 

ETYMOLOGY – The epithet refers to the similarities of the new species with T. zebrinum. 


Thallus moderately thin to moderately thick, epi- to hypophloedal, up to c. 400 µm high, 

partly bulging and flaking away from the substrate, pale grayish-green. Surface dull, smooth, 

rugose to verrucose, unfissured. Thallus covered by an incontinuous, thin protocortex up to c. 

15 µm thick. Algal layer well developed and continuous, calcium oxalate crystals abundant, 

predominantly large and scattered. Vegetative propagules not seen. Ascomata inconspicuous, 

moderately large, up to c. 600 µm in diam., roundish to irregular in fused ascomata, apo- to 

perithecioid, erumpent, regenerating, solitary to marginally fused, immersed. Disc sometimes 

becoming partly visible, whitish, epruinose. Pores small to moderately wide, up to c. 350 µm 

in diam., roundish to angular, formed by apical margin of proper exciple, pore margin/visible 

part of proper exciple split, off-white to dark-brown or blackish, incurved. Thalline rim 

moderately thick, becoming ±distinctly layered with proceeding succession of ascomata 

generations, split to lacerate, dark brownish to blackish, sometimes indistinctly contrasted by 

off-white to pale brownish layers, outer thalline rim concolorous with thallus, in distinctly 

layered ascomata thalline rim somewhat exfoliating, incurved in inner layer(s) to erect in 

outer layer(s), in un-layered ascomata thalline rim predominantly incurved. Exciple free in 

apical parts, (moderately) thick, yellowish-brown internally, distinctly carbonized marginally, 

non-amyloid. Hymenium up to c. 120 µm high, non-inspersed, strongly conglutinated, 

paraphyses straight, parallel, unbranched, with unthickened tips, lateral paraphyses present, 

short but usually conspicuous, up to c. 15 µm long, columellar structures absent, 

subhymenium ±distinctly carbonized. Epihymenium indistinct to thin, hyaline, sometimes 

with fine grayish granules. Asci 8-spored, tholus moderately thick, becoming moderately thin


with maturity. Ascospores (moderately) small, transversely septate in younger stages to 

±submuriform at maturity, cell walls (moderately) thin, endospore moderately thick, with thin 

halo, hyaline, non-amyloid, fusiform to claviform with roundish to subacute ends, loci 

roundish, oblong to lentiform, end cells hemispherical to conical, transverse septae thin, 

regular, 25-35 x 7-10 µm with 10-14 x 1-2(3) loci. Pycnidia not seen. 


Fig. 209. Thelotrema subzebrinum: growth habit (A), ascomata (B, C) and ascospores (D-F). A.-F.: 

CANB-holotype. Bar= A: 1 mm; B, C: 0.5 mm; D: 7 µm; E: 5 µm; F: 6 µm.


ECOLOGY AND DISTRIBUTION – Thelotrema subzebrinum 

grows on bark in a warm-temperate 

rainforest at 850 m. It is rare and only known from 

northern New South Wales. 

NOTES – The new species is characterized by a 

thick, ecorticate thallus, ascomata of the L. schizoloma-type, 

moderately small, transversely septate 

to submuriform, hyaline, non-amyloid ascospores 

and the absence of secondary compounds. It is 

similar to T. zebrinum, which differs by larger (up 

to 80 µm long with up to 22 loci), strictly 

transversely septate ascospores and the presence of 

the protocetraric and/or stictic acid chemosyndromes. 

Thelotrema parvizebrinum is readily 

distinguished by smaller ascospores (up to 20 µm 

long, with up to 8 loci). 


“Thelotrema” zebrinum Mangold spec. nov. ined. 

Type: Australia, New South Wales, Mt. Warning NP., track from summit to parking lot, Mangold 19 zo 

(CANB-holotype; NSW-isotype). 

ETYMOLOGY – The epithet refers to the layered ascomata margins (from lat. zebrinus 

=striped). 



T. subzebrinum. 

Thallus variable, thin to very thin to partly entirely hypophloedal, up to c. 100 µm high, 

rarely pale yellowish-gray or greenish-olive to more often pale gray to grayish-green. Surface 

dull to shiny, smooth to roughened, continuous to verruculose, sometimes fissured. Thallus 

predominantly covered by an incontinuous protocortex, up to c. 20 µm thick, rarely becoming 

distinctly conglutinated forming a true cortex of periclinal hyphae. Algal layer incontinuous 

and poorly developed, calcium oxalate crystals sparse, ±small, solitary or in clusters. 

Vegetative propagules not seen. Ascomata variable, usually conspicuous, (moderately) large, 

up to c. 0.9 mm in diam., roundish, peri- to indistinctly apothecioid in older stages, erumpent, 

solitary, regenerating, predominantly ±emergent, (irregular-)hemispherical to (irregular- 

)depressed-urceolate. Disc usually not visible from surface, rarely becoming partly visible, 

pale grayish to whitish, epruinose. Pores tiny to (moderately) small, up to c. 200 µm in diam., 

roundish to irregular, formed by apical margin of proper exciple, pore margin/visible part of 

proper exciple ±split, fused to indistinctly free, off-white to pale brownish, incurved. Thalline 

rim (moderately) thick, becoming distinctly layered with proceeding succession of ascomata 

generations, coarsely split to predominantly lacerate, sometimes eroded, (dark) brownish to 

blackish, ±distinctly contrasted by off-white layers, outer thalline rim concolorous with 

thallus, in distinctly layered thalline rims exfoliating, incurved to erect in inner layer(s), erect 

to slightly recurved in outer layer(s), in un-layered thalline rims predominantly incurved. 

Proper exciple moderately thick, predominantly apically free, rarely with a very thin, 

indistinct hyaline area internally to ±entirely (dark-)brown or usually distinctly carbonized, 

sometimes with substrate inclusions, distinctly amyloid at the base. Inner subhymenial layer


dark-brown to carbonized, sometimes followed by a hyaline layer of newly developing 

hymenium. Hymenium up to c. 200 µm high, non-inspersed, strongly conglutinated, 

paraphyses straight to slightly bent, ±distinctly parallel, unbranched, tips unthickened, lateral 

paraphyses present, conspicuous, up to c. 30 µm long, true columella absent, columella-like 

structures sometimes present in newly developing ascomata generations (=raised subhymenial 

layer). Epihymenium indistinct, hyaline, without granules or crystals. Asci 6-8-spored, tholus 

and lateral ascus walls (moderately) thick, thin when mature. Ascospores predominantly 

moderately large, transversely septate, cell walls (moderately) thick, becoming distinctly 

crenate at maturity, with thin and somewhat irregular halo in younger stages, non-halonate at 

maturity, hyaline, non-amyloid, oblong to oblong-ellipsoid, often slightly bent, with rounded 

to narrowed-rounded ends, loci large, roundish to slightly angular (predominantly in younger 

ascospores), oblong to usually ±lentiform, end cells hemispherical to conical, septae 

(moderately) thin, predominantly regular, 30-80 x 6-11 µm with 12-22 loci. Pycnidia not 

seen. 

CHEMISTRY – Thallus K+ yellowish to yellow, C-, PD+ reddish to orange; containing 

constictic, conprotocetraric, stictic, protocetraric, fumarprotocetraric (major to absent), 

hypoconstictic, hypostictic and virensic (minor to absent) acids. 

Fig. 211. Thelotrema zebrinum: growth habit (A), ascomata (B, C), ascoma section (D) and 

ascospores (E, F). A., B., E., F.: CANB-holotype; C., D.: Wilson s.n. (NSW-169660). Bar= A: 

1.75 mm; B, C: 0.5 mm; D: 200 µm; E: 22 µm; F: 7 µm.



zebrinum occurs on tree bark in cool- to warmtemperate 

or subtropical rainforests in altitudes 

ranging from 250 to 1200 m. It occurs in the 

Queensland/New South Wales-border-region, Lord 

Howe Island (NSW), southern Victoria and 

Tasmania. It is an Australasian species that is also 

known from New Zealand (Lumbsch & al. 2008). 

NOTES – The new species is readily characterized 

by the layered and ±distinctly carbonized 

ascoma margins, the presence of lateral paraphyses 

and moderately large, trans-versely septate, thickwalled, 

hyaline, non-amyloid ascospores. The 

secondary metabolites found are remarkable, 

several collections have either substances of the 

stictic or the protocetraric acid chemosyndromes, 

however, a few specimen produce acids of both 

chemosyndromes. Consequently, it is refrained 

from separating two chemical strains for this taxon. 


T. zebrinum. 

Thelotrema guadeloupensis and Leptotrema schizoloma are similar, but both have 

eumuriform ascospores and can be therefore easily separated. For differences to T. 

parvizebrinum and T. subzebrinum see under these species. Müller (1893) identified Wilson’s 

collection from Victoria (NSW-Wilson 949) erroneously as “Ocellularia” gyrostomoides, an 

unidentified Stictis species from Queensland. Ocellularia concentrica from New Zealand 

transferred from Odontotrema by Sherwood (1987) might be an older name for T. zebrinum. 

The type, however, was not available for study. 

SPECIMENS EXAMINED – Australia, Queensland, Gambubal SF., E of Emu Vale, Hafellner & Stevens 16395 

(GZU). New South Wales, Lord Howe Island, Goat House Cave, Elix 42267 pr.p. (CANB, [B-dubl. is a different 

species]). Victoria: Eastern Highlands, Black Spur: Nov. 1900, Wilson s.n. (MEL-724545); Feb. 1888, Wilson 

s.n. (NSW-539404); Jan. 1890, Wilson s.n. (NSW-539406). Warburton, Wilson 949 (NSW). Ottway NP., 10 km 

W of Apollo Bay, Mangold 1 g (F). Tasmania: Ulverstone, Wilson s.n. (NSW-539324). Mt. Arthur, Feb. 1891, 

Wilson s.n. (BM-761740, NSW-539403). 6 km S of Waratah Hwy. jct. with the Murchison Hwy., Hale 68737, 

68738 (US). Walls of Jerusalem NP., Kantvilas 65/99 (HOB). Mt. Wellington: Wilson s.n. (NSW-169662); 

Brown Track, Wilson s.n. (NSW-153763, -169660); St. Crispin's Well, Wilson s.n. (NSW-169661).

3. Molecular phylogeny 294 

3. Molecular phylogeny 

3. 1. Introduction 

Thus far, molecular studies on Graphidaceae s. lat. are rather limited. One sequence was 

included in an early study addressing the circumscription of Ostropales (Winka & al., 1998), 

while Kauff & Lutzoni (2002) included three species in their study on the phylogenetic 

relationships of Gyalectales and Ostropales. In studies addressing phylogenetic relationships 

of other families related to Ostropales, several sequences of thelotremataceaen Graphidaceae 

were included by Grube & al. (2004) and Lücking & al., (2004), a study by Martín & al., 

(2003) dealt with the molecular phylogeny of the genus Diploschistes. A molecular study 

with special emphasis on the systematic position of Nadvornikia was carried out by Lumbsch 

& al. (2004a). In an analysis addressing the phylogeny of Lecanoromycetes, Miadlikowska & 

al. (2006) included six species of two genera of thelotrematoid Graphidaceae using five gene 

regions. As the most recent treatments exclusively focussed on Graphidaceae s. lat., a singlegene 

phylogenetic analyses was performed on genera of thelotremataceaen Graphidaceae 

(Frisch & al., 2006), and a two-gene study addressing Graphidaceae s. str. (Staiger & al., 

2006). 

As already implied above, molecular phylogenetic studies are based on various genes and 

DNA regions. The most common genes used in fungal systematics are mitochondrial and 

nuclear ribosomal DNA (mt rDNA/nu rDNA). In previous studies it could be shown 

(Lumbsch & al. 2000, 2001a, Schmitt & al. 2001) that the most appropriate genetic markers 

for phylogenetic tests on the family/generic level are the large subunit (28s, LSU) encoding 

gene of the nu rDNA and the small subunit (18s, SSU) encoding gene of the mt rDNA. A 

combined dataset of both genes was used for the analysis focusing on the family-level 

phylogeny (analysis 1), for the molecular revision of the phylogeny of Thelotrema and 

Topeliopsis, single gene data sets were employed, using nu LSU rDNA for Topeliopsis 

(analysis 2) and mt SSU rDNA for Thelotrema (analysis 3). 

3. 2. Material and methods 

3. 2. 1. Material 

The examined material was predominantly collected on the field trips to Pacific Australia 

(see chapter 2. 4. 1.). In addition, herbarium specimens from BG, CANB, F, OTA, and PH 

were used. Several previously published sequences of mt SSU rDNA and nu LSU rDNA, 

available from GenBank were also included. 

For analysis 1, data matrices of 105 ascomycetes were assembled. Nine taxa of Ostropales, 

which do not belong to Graphidaceae, were included as outgroup. For analysis 2, sequence 

data of 40 specimens of 38 species were assembled. Myriotrema desquamans and Thelotrema 

glaucopallens were used as outgroup, since Myriotrema and the T. glaucopallens-group 

clustered basal to the ingroup taxa in analysis 1. For analysis 3, data matrices of 43 sequences 

of 19 Thelotrema species were assembled. As an outgroup, seven sequences of six Chapsa 

species were included since the genus was sister-group to Thelotrema s. str. in analysis 2. In 

the tables 5-7. all used species and specimen are listed with further specifications.


Table 5: Analysis 1. Species and specimens used in the current study with GenBank accession numbers. 

Species Sample 

GenBank acc. no. 

mt SSU nu LSU 

"Graphina" peplophora DQ431966 DQ431930 

"Sarcographina" glyphiza DQ431972 DQ431934 

"Thelotrema" zebrinum Australia, New South Wales, Mangold 19 zo (F) EU075608 EU075652 

Absconditella sp. AY300873 AY300825 

Absconditella sphagnorum AY300872 AY300824 

Acanthothecis aurantiaca DQ431965 DQ431929 

Acanthotrema brasilianum DQ384916 DQ431928 

Acarosporina microspora AY584612 AY584643 

Ampliotrema auratum 1 Australia, Queensland, Lumbsch 19160 wA & Mangold (F) EU075564 EU075612 

Ampliotrema auratum 2 Australia, Queensland, Mangold 33a (F) EU075565 EU075613 

Chapsa astroidea Australia, Queensland, Lumbsch 19166 n & Mangold (F) EU075566 EU075614 

Chapsa cf. pulchra] Australia, Queensland, Lumbsch 19082 e & Mangold (F) EU075570 EU075618 

Chapsa leprocarpa Australia, Queensland, Lumbsch 19125 k & Mangold (F) EU075568 EU075615 

Chapsa niveocarpa Australia, Queensland, Lumbsch 19151 p & Mangold (F) EU075567 EU075616 

Chapsa phlyctidioides Australia, Queensland, Lumbsch 19100 f & Mangold (F) EU075569 EU075617 

Chapsa pulchra Australia, Queensland, Lumbsch 19129 t & Mangold (F) EU075571 EU075619 

Chroodiscus coccineus DQ384915 AF465441 

Cryptodiscus foveolaris AY661673 AY661683 

Cryptolechia sp. . Kenya, Western Province, Lumbsch 19562 g, Mangold & 

Divakar (F) 

EU075572 EU075620 

Diorygma circumfusum DQ431963 AY640019 

Diorygma junghuhnii DQ431962 AY640018 

Diorygma pruinosum DQ431964 AY640014 

Diorygma sipmanii DQ431961 AY640020 

Diploschistes 

cincereocaesius 

AY300885 AY300835 

Diploschistes muscorum AY300886 AY300836 

Diploschistes rampoddensis AF431954 AF274094 

Diploschistes thunbergianus AF431955 AF274095 

Dyplolabia afzelii DQ431949 DQ43192 

Fibrillithecis halei Australia, Queensland, Mangold 31g (F) EU075573 EU075621 

Fissurina marginata DQ431951 AY640012 

Fissurina triticea DQ431952 AY640011 

Glyphis cicatricosa DQ431955 AY640025 

Glyphis scyphulifera DQ431956 AY640027 

Glyphis substriatula DQ431982 AY640026 

Graphis caesiella DQ431975 AY640028 

Graphis chrysocarpa DQ431987 AF465448 

Graphis cinerea DQ431988 DQ431947 

Graphis cleistoblephara DQ431974 AY640031 

Graphis pavoniana DQ431986 DQ431946 

Graphis ruiziana DQ431985 DQ431945 

Graphis scripta AY853370 AY853322 

Hemithecium implicatum DQ431978 DQ431939 

Leiorreuma hypomelaenum DQ431971 DQ431933 

Leptotrema wightii Costa Rica, Lücking 2034A (F) EU075574 EU075622 

Leucodecton 

subcompunctum 1 

Australia, Queensland, Lumbsch 19116 o & Mangold (F) EU075575 EU075623 

Leucodecton 

subcompunctum 2 

Australia, Queensland, Lumbsch 19153 p & Mangold (F) EU075576 EU075624 

Myriotrema microporum Australia, Queensland, Lumbsch 19092 o & Mangold (F) EU075578 EU075626 

Myriotrema olivaceum 1 DQ384900 AY640009 

Myriotrema olivaceum 2 Australia, Queensland, Lumbsch 19113f & Mangold (F) EU075579 EU075627




mt SSU nu LSU 

Myriotrema trypaneoides Australia, Queensland, Lumbsch 19167 v & Mangold (F) EU075580 EU075628 

Nadvornikia hawaiensis Australia, Queensland, Mangold 36 w (F) EU075581 AY605080 

Neobelonia sp. AY300879 AY300830 

Ocellularia albocincta Costa Rica, Lücking 063 (F) EU075585 EU075632 

Ocellularia cavata DQ384877 DQ431935 

Ocellularia chiriquiensis Australia, Queensland, Mangold 18 d (F) EU075582 EU075629 

Ocellularia diacida Australia, Queensland, Lumbsch 19120 jb & Mangold (F) EU075583 EU075630 

Ocellularia inturgescens Australia, Queensland, Lumbsch 19132 w & Mangold (F) EU075577 EU075625 

Ocellularia massalongoi Australia, Queensland, Mangold 36 m (F) EU075584 EU075631 

Ocellularia papillata Australia, Queensland, Mangold 43 o (F) EU075586 EU075633 

Ocellularia perforata 1 DQ384881 AY605081 

Ocellularia perforata 2 Australia, Queensland, Lumbsch 19120 ja & Mangold (F) EU075587 EU075634 

Ocellularia postposita DQ384873 AY640008 

Ocellularia profunda 1 Australia, Queensland, Lumbsch 19116c & Mangold (F) EU075588 EU075635 

Ocellularia profunda 2 Australia, Queensland, Lumbsch19100p & Mangold (F) EU075589 NEW 

Ocellularia profunda 3 Australia, Queensland, Lumbsch 19123 k & Mangold (F) EU075590 EU075636 

Ocellularia sp. Australia, Queensland, Lumbsch 19144d & Mangold (F) EU075591 EU075637 

Ocellularia thelotremoides Australia, Queensland, Lumbsch 19108 I & Mangold (F) EU075592 EU075638 

Odontotrema sp. 1 AY661674 AY661684 

Odontotrema sp. 2 AY661675 AY661685 

Phaeographis brasiliensis DQ431958 AY640022 

Phaeographis caesioradians DQ431968 AY640021 

Phaeographis intricans DQ431960 DQ431927 

Phaeographis lecanographa DQ431983 DQ431943 

Phaeographis lobata DQ431984 DQ431944 

Platygramme australiensis DQ431970 AY640024 

Platygramme 

caesiopruinosa 1 

DQ431973 AY640023 

Platygramme 

caesiopruinosa 2 

Australia, Queensland, Mangold 30 eI (F) EU075593 EU075639 

Ramonia sp. AY300921 AY300871 

Sagiolechia rhexoblephara AY853341 AY853391 

Sarcographa fennicis DQ431967 DQ431931 

Sarcographa ramificans DQ431981 DQ431942 

Stegobolus cf. subcavatus Australia, Queensland, Lumbsch 19151 u & Mangold (F) EU075595 EU075641 

Stegobolus fissus Australia, Queensland, Lumbsch 19108f & Mangold (F) EU075594 EU075640 

Stictis populorum AY300882 AY300833 

Stictis radiata AY300914 AY300864 

Thelotrema bicinctulum Australia, Queensland, Mangold 34 f (F) EU075598 EU075642 

Thelotrema cf. nureliyum Australia, Queensland, Lumbsch 19110 b & Mangold (F) EU075603 EU075648 

Thelotrema crespoae Australia, New South Wales, Mangold 27 v (F) EU075606 NEW 

Thelotrema diplotrema Australia, Queensland, Lumbsch 19127 v & Mangold (F) EU075599 EU075643 

Thelotrema gallowayanum Australia, Queensland, Lumbsch 19151 k & Mangold (F) EU075600 EU075653 

Thelotrema glaucopallens DQ384906 AY605069 

Thelotrema lepadinum AY300916 AY300866 

Thelotrema monosporum 1 Australia, Queensland, Lumbsch19161 xb & Mangold (F) EU075596 EU075644 

Thelotrema monosporum 2 Australia, Queensland, Lumbsch19158 w & Mangold (F) EU075601 EU075646 

Thelotrema nureliyum 1 Australia, Queensland, Lumbsch 19100 ya & Mangold (F) EU075597 EU075647 

Thelotrema nureliyum 2 Australia, New South Wales, Mangold 22c (F) EU075604 EU075649 

Thelotrema pachysporum DQ384918 DQ431925 

Thelotrema rugatulum Australia, Queensland, Lumbsch 19082b & Mangold (F) EU075605 EU075650 

Thelotrema saxatile USA, Maryland, Lendemer 6389 (PH) EU075602 EU075645 

Thelotrema subtile 1 DQ871020 DQ871013 

Thelotrema subtile 2 Australia, Victoria, Mangold 3j (F) EU075607 EU075651 

Thelotrema suecicum AY300917 AY300867




mt SSU nu LSU 

Topeliopsis decorticans Australia, Victoria, Mangold 5i.I (F) EU075609 EU075654 

Topeliopsis meridensis Costa Rica, Lücking 17770 (F) EU075610 EU075655 

Topeliopsis muscigena Australia, Vicotria, Mangold 5d (F) EU075611 EU075656 

Table 6: Analysis 2. Species and specimens used in the current study with GenBank accession numbers. 

Species Sample GenBank 

acc. no. 

nu LSU 

"Thelotrema" zebrinum Australia, New South Wales, Mangold 19 zo (F) EU075652 

Acanthotrema brasilianum DQ431928 

Chapsa astroidea Australia, Queensland, Lumbsch 19166 n & Mangold (F) EU075614 

Chapsa phlyctidioides Australia, Queensland, Lumbsch 19100 f & Mangold (F) EU075617 

Chapsa pulchra Australia, Queensland, Lumbsch 19129 t & Mangold (F) EU075619 

Chroodiscus coccineus AF465441 

Diploschistes cincereocaesius AY300835 

Diploschistes diploschistoides AY605076 

Diploschistes elixii Australia, Western Australia, Elix 32450 (F, isotype) EU126644 

Diploschistes muscorum AY300836 

Diploschistes rampoddensis AF274094 

Diploschistes scruposus AF279389 

Diploschistes thunbergianus AF274095 

Melanotopelia toensbergii USA, Washington, Tønsberg 32310 (BG) EU126657 

Myriotrema cinereum AY605074 

Myriotrema desquamans AY605085 

Myriotrema laeviusculum AY605070 

Myriotrema olivaceum Australia, Queensland, Lumbsch 19092 g & Mangold (F) EU126645 

Ocellularia bahiana AY605067 

Ocellularia bonplandii Australia, Queensland, Lumbsch 19092 f & Mangold (F) EU126646 

Ocellularia kalbii Australia, Queensland, Lumbsch 19085 h & Mangold (F) EU126647 

Ocellularia microstoma Australia, Queensland, Lumbsch 19108 c & Mangold (F) EU126648 

Ocellularia postposita AY640008 

Ocellularia thelotremoides Australia, Queensland, Lumbsch 19081 j & Mangold (F) EU126649 

Thelotrema diplotrema Australia, Queensland, Lumbsch 19127 v & Mangold (F) EU075643 

Thelotrema glaucopallens AY605069 

Thelotrema lepadinum AY300866 

Thelotrema myriocarpum Australia, Queensland, Lumbsch 19120 d & Mangold (F) EU126650 

Thelotrema pachysporum DQ431925 

Thelotrema porinaceum Australia, Queensland, Lumbsch 19108 d & Mangold (F) EU126651 

Thelotrema porinoides Australia, Queensland, Lumbsch 19113 s & Mangold (F) EU126652 

Thelotrema suecicum AY300867 

Thelotrema weberi AY605084 

Topeliopsis acutispora Australia, New South Wales, Mangold 13 f (F) EU126653 

Topeliopsis decorticans Australia, Victoria, Mangold 5i.I (F) EU075654 

Topeliopsis meridensis Costa Rica, Lücking 17770 (F) EU075655 

Topeliopsis muscigena 1 Australia, Victoria, Mangold 5iII (F) EU126654 

Topeliopsis muscigena 2 Australia, Vicotria, Mangold 5d (F) EU126655 

Topeliopsis muscigena 3 EU075656 

Topeliopsis subdenticulata New Zealand, Otago, 25.4.2006, Knight (OTA - 59967) EU126656


Table 7: Analysis 3: Species and specimens used in the current study with GenBank accession numbers. 

Species Sample GenBank 

acc. no. 

mt SSU 

Chapsa astroidea Australia, Queensland, Lumbsch 19116 n & Mangold (F) AY300917 

Chapsa indica DQ384911 

Chapsa leprocarpa Australia, Queensland, Lumbsch 19125 k & Mangold (F) EU075568 

Chapsa niveocarpa Australia, Queensland, Lumbsch 19151 p & Mangold (F) EU075567 

Chapsa phlyctidioides 1 Australia, Queensland, Lumbsch 19100 f & Mangold (F) EU075569 

Chapsa phlyctidioides 2 Australia, Queensland, Mangold 39 ze (F) NEW 

Chapsa pulchra Australia, Queensland, Lumbsch 19129 t & Mangold (F) EU075571 

Thelotrema capetribulense 1 Australia, Queensland, Lumbsch 19160 q & Mangold (F) NEW 

Thelotrema capetribulense 2 Australia, Queensland, Lumbsch 19161 xA & Mangold (F) NEW 

Thelotrema capetribulense 3 Australia, Queensland, Lumbsch 19162 I & Mangold (F) NEW 

Thelotrema conveniens Australia, Queensland, Mangold 29af (F) NEW 

Thelotrema crespoae Australia, New South Wales, Mangold 27 v (F) EU075606 

Thelotrema defossum Australia, Queensland, Lumbsch 19161 v & Mangold (F) NEW 

Thelotrema diplotrema 1 Australia, Queensland, Lumbsch 19127 v & Mangold (F) EU075599 

Thelotrema diplotrema 2 Australia, Queensland, Lumbsch 19161 r & Mangold (F) NEW 

Thelotrema diplotrema 3 Australia, Queensland, Lumbsch 19139 r & Mangold (F) NEW 

Thelotrema gallowayanum Australia, Queensland, Lumbsch 19151 k & Mangold (F) EU075600 

Thelotrema lepadinum 1 DQ431957 




Thelotrema lepadinum 5 Australia, Victoria, Mangold 1 a (F) NEW 

Thelotrema lepadinum 6 AY300916 

Thelotrema lepadodes Australia, Queensland, Lumbsch 19158 k & Mangold (F) NEW 

Thelotrema monosporum 1 Australia, Queensland, Lumbsch19158 w & Mangold (F) EU075601 

Thelotrema monosporum 2 Australia, Queensland, Lumbsch 19158 v & Mangold (F) NEW 

Thelotrema monosporum 3 - EU075596 

Thelotrema monosporum 4 DQ384919 

Thelotrema nureliyum 1 Australia, Queensland, Lumbsch19174 x & Mangold (F) NEW 

Thelotrema nureliyum 2 Australia, New South Wales, Mangold 22c (F) EU075604 

Thelotrema nureliyum 3 Australia, Queensland, Lumbsch 19174 I & Mangold (F) NEW 

Thelotrema nureliyum 4 Australia, Queensland, Lumbsch 19100 ya & Mangold (F) EU075597 

Thelotrema nureliyum 5 Australia, Queensland, Lumbsch19100 k & Mangold (F) NEW 

Thelotrema nureliyum 6 Australia, Queensland, Lumbsch 19110 b & Mangold (F) EU075603 

Thelotrema oleosum Australia, New South Wales, Mangold 25 m (F) NEW 

Thelotrema pachysporum 1 DQ384918 

Thelotrema pachysporum 2 Australia, Queensland, Lumbsch 19162 j & Mangold (F) NEW 

Thelotrema porinaceum Australia, Queensland, Lumbsch 19156 d & Mangold (F) NEW 

Thelotrema porinoides 1 DQ384920 

Thelotrema porinoides 2 Australia, Queensland, Lumbsch 19137 i & Mangold (F) NEW 

Thelotrema pseudosubtile Australia, Queensland, Lumbsch 19117 k & Mangold (F) NEW 

Thelotrema rugatulum Australia, Queensland, Lumbsch 19082b & Mangold (F) EU075605 

Thelotrema saxatile 1 Australia, Queensland, Lumbsch 19104 a & Mangold (F) NEW 

Thelotrema saxatile 2 U.S.A., Lendemer 2149 (PH) NEW 

Thelotrema saxatile 3 Australia, Queensland, Mangold 32 v (F) NEW 

Thelotrema subtile 1 DQ871020 

Thelotrema subtile 2 Australia, Victoria, Mangold 3j (F) EU075607 

Thelotrema subtile 3 Australia, Victoria, Mangold 3 e (F) NEW 

Thelotrema suecicum 1 AY300917 

Thelotrema suecicum 2 Australia, Victoria, Mangold 5 f (F) NEW


3. 2. 2. DNA extraction 

For DNA extraction, small pieces of samples were taken, usually ascomata. As a first step, 

samples were soaked in acetone for1 h to remove secondary metabolites. The acetone was 

then discarded and subsequently air-dried samples were crushed using plastic pistils after 

cooling with liquid nitrogen and then processed using the DNeasy Plant Mini Kit (Qiagen) or 

E.Z.N.A. Fungal MiniPrep Kit (Omega-Biotech) following the instructions of the 

manufacturer. 

3. 2. 3. PCR amplification and purification of PCR products 

PCR amplifications were made using Ready-to-Go-PCR Beads (Amersham-Biosciences) 

as described in Winka & al., (1998), or performed as described in the following. 25 µL PCR 

reactions contained: 2.5 µL buffer, 2.5 µL dNTP mix, 1 µL of each primer (10 µM), 5 µL 

BSA, 2 µL Taq, 2 µL genomic DNA extract and 9 µL distilled water. The genomic DNA 

extract was prepared of dilutions (10 -1 up to 10 -2 ) or undiluted DNA. Primers for 

amplification were: a) for the nu LSU rDNA: nu-LSU-0155-5' (Döring & al., 2000), nu-LSU- 

0042-5’ (=LR0R), nu-LSU-1432-3' (=LR7), and nu-LSU-1125-3' (=LR6) (Vilgalys & Hester, 

1990), and AL2R: GCGAGTGAAGCGGCAACAGCTC; b) for the mt SSU rDNA: mr SSU1 

(Zoller & al., 1999) and MSU 7 (Zhou & al., 2001). 

Thermal cycling parameters were: initial denaturation for 3 min at 94-95ºC, followed by 30 

cycles of 1 min at 95ºC, 1 min at 52ºC, 1 min at 73ºC, and a final elongation for 7 min at 

73ºC; or when PCR Beads were used, a) for nu LSU: initial denaturation for 5 min at 94ºC, 

followed by 35 cycles of 30 sec at 94ºC, 30 sec at 52ºC, 1 min at 72ºC, and a final elongation 

for 10 min at 72ºC, and b) for mt SSU: initial denaturation for 5 min at 94ºC, followed by 35 

cycles of 45 sec at 94ºC, 1 min at 50ºC, 1 min 30 sec at 72ºC, and a final elongation for 10 

min at 72ºC. 

Amplification products were viewed on 1% or 2% agarose gels stained with ethidium 

bromide and subsequently purified using the QIAquick PCR Purification Kit (Qiagen), 

Nucleo Spin DNA purification kit (Macherey-Nagel) or EZNA Cycle-Pure kit 200 (Omega 

Biotech). 

3. 2. 4. Sequencing and sequence assembly 

For sequencing the same sets of primers were used as for the PCR amplification (see 

above). Cycle sequencing was executed with the following program: 25 cycles of 95ºC for 30 

sec, 48ºC for 15 sec, and 60 º C for 4 min. If direct sequencing failed due to the presence of 

multiple products, PCR products were cloned using the TOPO TA cloning kit (Invitrogen) or 

pGEM-T easy vector II cloning kit (Omega Biotech). We picked 3–10 clones of each cloning 

reaction. Cloned products were sequenced with universal primers specific to the plasmids: 

M13for and M13rev (TOPO TA) and T7 and SP6 (pGEM-T). Sequenced products were 

precipitated with 10 µL of sterile dH2O, 2 µL of 3 M NaOAc, and 50 µL of 95% EtOH before 

they were loaded on an ABI 3100 or 3730 (Applied Biosystems) automatic sequencer. 

Sequence fragments obtained were assembled with SeqMan 4.03 (DNASTAR) and 

manually adjusted. The identity of the obtained consensus sequences was checked using the 

BLAST program (Altschul & al., 1990) in the GenBank database.


3. 2. 5. Sequence alignments 

Alignments for the single gene data sets were done using Clustal W (Thompson & al., 

1994). Insertion and ambiguously aligned regions were delimited manually. Since the 

mitochondrial data set contains sequence portions that are highly variable, standard multiple 

alignment programs, such as Clustal (ibid.) become less reliable when sequences show a high 

degree of divergence. Therefore, for the combined dataset of analysis 1 an alignment 

procedure that uses a linear Hidden Markov Model (HMM) as implemented in the software 

SAM (Sequence Alignment and Modelling system) (Karplus & al., 1998) was employed for 

separate alignments of the two data sets. Regions that were not aligned with statistical 

confidence were excluded from the phylogenetic analysis. 

3. 2. 6. Phylogenetic analysis 

The alignments were analysed by maximum parsimony (MP) and a Bayesian approach 

(B/MCMC) (Huelsenbeck & al., 2001; Larget & Simon, 1999). To test for potential conflict, 

parsimony bootstrap analyses were performed on each individual data set, and 75% bootstrap 

consensus trees were examined for conflict (Lutzoni & al., 2004). 

Maximum parsimony analyses were performed using the program PAUP* (Swofford, 

2003). Heuristic searches with 200 (analysis 1) and 1000 (analyses 2 and 3) random taxon 

addition replicates were conducted with TBR branch swapping and MulTrees option in effect, 

equally weighted characters and gaps treated as missing data. Bootstrapping (Felsenstein, 

1985) was performed based on 2000 replicates with random sequence additions. To assess 

homoplasy levels, consistency index (CI) and retention index (RI) were calculated from each 

parsimony search. 

The B/MCMC analyses were conducted using the MrBayes 3.1.1 program (Huelsenbeck & 

Ronquist, 2001). The analyses were performed assuming the general time reversible model of 

nucleotide substitution (Rodriguez & al., 1990), including estimation of invariant sites and 

assuming a discrete gamma distribution with six rate categories (GTR+I+G). In analysis 1, the 

data set was portioned into the two parts (nu LSU, mt SSU) and each partition was allowed to 

have own parameters as suggested by Nylander & al., (2004). No molecular clock was 

assumed. A run with 4,000,000 (analysis 1) and 20,000,000 (analyses 2 and 3) generations 

starting with a random tree and employing 12 (four in analysis 3) simultaneous chains was 

executed. Every 100 th tree was saved into a file. The first 200,000 (analysis 1), 800,000 

(analysis 2) and 1,000,000 (analysis 3) generations (i.e. the first 2000, 8000 or 10,000 trees 

respectively) were deleted as the "burn in" of the chain. The log-likelihood scores of sample 

points were plotted against generation time using TRACER 1.0 

(http://evolve.zoo.ox.ac.uk/software.html?id=tracer) to ensure that stationarity was achieved 

after the first 200,000, 800,000 and 1,000,000 generations respectively, by checking whether 

the log-likelihood values of the sample points reached a stable equilibrium value 

(Huelsenbeck & Ronquist, 2001). Of the remaining 76,000 (analysis 1), 384,000 (analysis 2) 

and 380,000 (analysis 3) trees (38,000, 192,000 or 190,000 from each of the parallel runs 

respectively) a majority rule consensus tree with average branch lengths was calculated using 

the sumt option of MrBayes. Posterior probabilities were obtained for each clade. Only clades 

with bootstrap support equal or above 70% (analysis 1) or equal or above 75% (single gene 

analyses) under MP and posterior probabilities ≥ 0.95 (in all cases) were considered as 

strongly supported. Phylogenetic trees were visualized using the program Treeview 1.6.6 

(Page, 1996). 

In the combined analysis as well as in the nu LSU single gene analysis, the results were 

incongruent with the current classification. It was therefore tested whether the data were 

sufficient to reject alternative topologies. For the results of the analysis 1 phylogeny the


following two topological constraints were tested: 1. monophyly of Graphidaceae and 2. 

monophyly of Thelotremataceae. Three topological constraints were tested for the phylogeny 

of analysis 2: 1. monophyly of Topeliopsis s. str. + T. toensbergii, 2. monophyly of 

Topeliopsis s. str. + T. meridensis, and 3. monophyly of Topeliopsis s. lat. For the hypothesis 

testing two different methods were used: a. Shimodaira-Hasegawa (SH) test (2000) and b. 

expected likelihood weight (ELW) test following Strimmer & Rambaut (2002). The SH and 

ELW tests were performed using Tree-PUZZLE 5.2 (Schmidt & al., 2002) with the combined 

data set on a sample of 200 unique trees, the best trees agreeing with the null hypotheses, and 

the unconstrained ML tree. These trees were inferred in Tree-PUZZLE employing the 

GTR+I+G nucleotide substitution model. 

3. 3. Results 

3. 3. 1. Analysis 1 

A matrix of 105 sequences was produced with 1164 unambiguously aligned nucleotide 

position characters.. This included 638 positions of the nu LSU and 526 of the mt SSU data 

set. 516 characters in the alignment were constant. Insertions present in the nu LSU rDNA of 

many taxa were removed from the alignment. The MP 75% bootstrap support method for 

testing data sets for incongruence indicated no strongly supported conflict (data not shown) 

and hence a combined analysis was performed. The combined alignment will be available in 

TreeBASE (http://www.treebase.org/treebase). Maximum parsimony analysis yielded in 25 

most parsimonious trees 3861 steps long (CI=0.28, RI=0.61). 144 positions in the matrix were 

parsimony-uninformative, and 504 informative. 

In the B/MCMC analysis of the combined data set, the likelihood parameters in the sample 

had the following mean (Variance): LnL = -20254.95 (0.51), base frequencies ϖ(A){mtSSU} 

= 0.353 (0.0003), ϖ(C) {mtSSU} = 0.124 (0.0002), ϖ(G){mtSSU} = 0.222 (0.0002), 

ϖ(T){mtSSU} = 0.301 (0.0003), rϖ(A){nuLSU} = 0.252(0.0003), ϖ(C) {nuLSU} = 0.204 

(0.0002), ϖ(G){nuLSU} = 0.308 (0.0003), ϖ(T){nuLSU} = 0.235(0.0003), rate matrix 

r(AC){mtSSU} = 0.073 (0.001), r(AG){mtSSU} = 0.221 (0.0005), r(AT){mtSSU} = 0.102 

(0.002), r(CG){mtSSU} = 0.092 (0.0002), r(CT){mtSSU} = 0.442 (0.0006), r(GT){mtSSU} = 

0.072 (0.0001), rate matrix r(AC){nuLSU} = 0.07 (0.0001), r(AG){nuLSU} = 0.177 

(0.0004), r(AT){nuLSU} = 0.09 (0.002), r(CG){nuLSU} = 0.394 (0.0008), r(CT){nuLSU} = 

0.581 (0.005), r(GT){nuLSU} = 0.044 (0.0008), the gamma shape parameter alpha{mtSSU} 

= 0.56 (0.0009), alpha{nuLSU} = 0.731 (0.0005), and the proportion of invariable site 

p(invar){mtSSU} = 0.242 (0.0004), and p(invar){nuLSU} = 0.316 (0.0003). 

The topologies of the MP and B/MCMC analyses did show one strongly supported 

conflict. Thelotrema rugatulum clustered unsupported in the MP tree at the base of the T. 

lepadinum clade (=Thelotrema s. str.), while it is nested within Topeliopsis with a long branch 

in the B/MCMC tree. Since a long branch leads to T. rugatulum the placement of this taxon is 

regarded as uncertain. As no further conflicts were found, only the 50% majority-rule 

consensus tree of Bayesian tree sampling is shown. Those nodes that received strong support 

(i.e. PP ≥0.95 in B/MCMC analysis and MP bootstrap ≥70%) in both the MP and Bayesian 

analyses are in bold (Fig. 213). 

In this phylogeny, it can be seen, that the clade comprising Graphidaceae s. lat. is strongly 

supported. Taxa hitherto placed in the two distinct families Graphidaceae and 

Thelotremataceae do not form two separate clades. Although the backbone of the phylogeny 

within the Graphidaceae/Thelotremataceae clade lacks supports, several, well-supported 

clades can be distinguished within this clade. A basal clade (Fissurina-group A) including


Fig. 213. Phylogeny of Graphidaceae s. lat. as inferred from a two gene-partition analysis. Type species 

of ingroup genera indicated by an asterisk. Graphidaceae s. str. taxa are indicated by a black bar (right 

scale), a gray bar represents taxa of thelotrematoid Graphidaceae. Groups mentioned in the text are 

marked by letters. The bottom scale indicates the genetic distance represented by the branch lengths. 

(For further explanations see text.)


Dyplolabia and Fissurina (both Graphidaceae s. str.) is strongly supported and forms a sistergroup 

to the remaining taxa of Graphidaceae s. str. and thelotrematoid groups. This sistergroup 

relationship, however, lacks support. Another well-supported clade (Ocellulariagroup 

B) includes taxa currently classified in Ampliotrema, Fibrillithecis, Myriotrema s. str. 

(except M. trypaneoides), Ocellularia, and Stegobolus. This clade forms a sister-group to an 

unsupported clade including all remaining Graphidaceae s. lat. Both Myriotrema and 

Ocellularia as currently circumscribed are polyphyletic, but there is no backbone support in 

the Ocellularia-clade. Fibrillithecis is clustered within Myriotrema, but this lacks support. 

Both Ampliotrema and Stegobolus are nested within Ocellularia, but also here, support is 

lacking. 

The remaining taxa are scattered in several groups with little backbone support. The 

relationship of Acanthothecis (group C) is not resolved. Acanthotrema (group D) appears as 

sister to Diploschistes (group E), which is strongly supported as monophyletic, but the 

relationships with other genera remain open; three further unsupported clades, all including 

taxa of both Graphidaceae s. str. and thelotremataceaen Graphidaceae, also have unresolved 

relationships among each other. The first (Topeliopsis-group F) is a weakly supported 

assemblage of different taxa, including “Graphina” peplophora and the thelotrematoid taxa 

Leptotrema wightii, Nadvornikia hawaiensis, two species of Topeliopsis, Myriotrema 

trypaneoides, and four species of Thelotrema. However, support within this clade is mostly 

lacking. Another clade includes the bulk of graphidaceaen taxa, with the thelotrematoid 

Chroodiscus coccineus (group G) unsupported at the base. The genera Diorygma (group I), 

Glyphis (group J), Phaeographis (group K), and Platygramme (group L), are monophyletic, 

all except Phaeographis strongly supported. Leiorreuma hypomelaenum is nested within 

Sarcographa. This group (M) forms a well-supported sister-group relationship to 

Platygramme. Four species of Graphis form a well-supported group (H) sister to the 

remaining Graphidaceae s. str. 

In the upper part of the tree an unsupported group includes two strongly supported clades; 

one comprising of Graphis s. str. (Hemithecium implicatum nested within; group N), and the 

other composed of thelotrematoid taxa currently classified in four genera: Chapsa, 

Leucodecton, Thelotrema s.str., and one Topeliopsis species. Within this clade Chapsa is 

paraphyletic (group O) with species of the other three genera nested within. The relationships 

of Leucodecton subcompunctum and Topeliopsis meridensis, however, lack support. 

Thelotrema s. str. forms a well-supported monophyletic group (group P). 

The two alternative topology tests both significantly rejected monophyly of Graphidaceae 

and monophyly of Thelotremataceae as currently circumscribed (p


Fig. 214. Phylogeny of Topeliopsis and allied genera of thelotremataceaen Graphidaceae as inferred from 

a nu LSU rDNA data set. Groups mentioned in the text are marked by letters. The bottom scale 

indicates the genetic distance represented by the branch lengths. (For further explanations see text.)


Since the topologies of the MP and B/MCMC analyses did not show any strongly 

supported conflicts, only the 50% majority-rule consensus tree of Bayesian tree sampling is 

shown (Fig. 214). Those nodes that received strong support (i.e. PP ≥0.95 in B/MCMC 

analysis and MP bootstrap ≥75%) in both the MP and Bayesian analyses are in bold. 

In the majority-rule consensus tree, Topeliopsis is polyphyletic, with a core group, 

including the type species T. muscigena, and T. acutispora, T. decorticans and T. 

subdenticulata in a well-supported monophyletic clade (clade A), while T. meridensis and the 

former T. toensbergii (=Melanotopelia toensbergii) cluster in other groups. Topeliopsis 

meridensis, is basal to a well supported clade including Chapsa and Thelotrema s. str., 

however, its basal placement is unsupported. Melanotopelia toensbergii has a basal placement 

to a clade including the well-supported Chapsa-Thelotrema s. str.-T. meridensis clade and 

Chroodiscus coccineus (clade D). The relationships of these groups are strongly supported, 

but the exact position of M. toensbergii lacks support. The relationships of Topeliopsis s. str. 

are not resolved with confidence. The genus Diploschistes is strongly supported as 

monophyletic (clade B). As also shown in the other phylogenetic analyses (1 and 3), the 

genera Chapsa, Myriotrema, Ocellularia, and Thelotrema as currently circumscribed are not 

monophyletic. Several taxa of Ocellularia and two Myriotrema species cluster in a well 

supported clade (clade C) basal to clade D and an unresolved group of three different genera, 

including Thelotrema glaucopallens, Acanthotrema brasilianum and Myriotrema olivaceum. 

“Thelotrema” zebrinum, the only member of the Leptotrema schizoloma-group examined, is 

basal to the Topeliopsis s. str. clade, but its relationship to this clade as well as to the other 

taxa is unresolved. 

3. 3. 3. Analysis 3 

43 sequences were aligned to produce a matrix of 698 unambiguously aligned nucleotide 

position characters; 484 characters in the alignment were constant. Maximum parsimony 

analysis yielded 288 most parsimonious trees (438 steps long). The strict consensus tree did 

not contradict the Bayesian tree topology. 

In the B/MCMC analysis of the combined data set, the likelihood parameters in the sample 

had the following mean (Variance): LnL = -3552.639 (0.32), base frequencies ϖ(A) = 0.334 

(0.00081), ϖ(C) = 0. 157 (0.00076), ϖ(G) = 0.202 (0.00076), ϖ(T) = 0.308 (0.0008), the 

gamma shape parameter alpha = 0.609 (0.0017), and p(invar) = 0.371 (0.00052). 

Since the topologies of the MP and B/MCMC analyses did not show any strongly 

supported conflicts, only the 50% majority-rule consensus tree of Bayesian tree sampling is 

shown. Those nodes that received strong support (i.e. PP ≥0.95 in B/MCMC analysis and MP 

bootstrap ≥70%) in both the MP and Bayesian analyses are in bold as shown in Figure 215. 

Thelotrema is strongly supported as monophyletic in the majority-rule consensus tree. The 

backbone of the phylogeny within Thelotrema is not resolved with confidence, with the single 

exception of a placement of a group of five species (T. conveniens, T. crespoae, T. 

gallowayanum, T. oleosum, T. porinaceum) at the base. The relationships among these basal 

species are not clear, with the exception of a strongly supported sister-group relationship of T. 

oleosum and T. porinaceum (clade A). Within the group of remaining Thelotrema species, six 

well supported clades (clades B-G) can be distinguished; the relationships between these 

clades do not receive strong support and hence are uncertain. The new species T. 

capetribulense forms a well supported sister-group to T. porinoides (clade B). Three samples 

of T. diplotrema cluster together (clade C), but the relationships of this species remain 

unresolved, which is also true for the relationships of T. nureliyum (clade D) and T. subtile 

(clade F). Within clade E five species cluster together. Thelotrema lepadinum has a well 

supported sister-group relationship to T. suecicum and these two species are strongly 

supported as sister to T. rugatulum, which are sister to an unsupported sister-group, consisting


Fig. 215. Phylogeny of Thelotrema and Chapsa (outgroup) as inferred from a mt SSU rDNA data set. 

Groups mentioned in the text are marked by letters. The bottom scale indicates the genetic distance 

represented by the branch lengths. (For further explanations see text.)


of T. defossum and T. pseudosubtile. Clade G includes four species, T. saxatile forms a well 

supported sister-group with T. lepadodes, while T. monosporum is nested within T. 

pachysporum, but this position lacks support. The three samples of T. saxatile cluster 

together, but the relationships of this species within clade G remain uncertain. 

3. 4. Discussion 

3. 4. 1. Phylogenetic relationship of Graphidaceae and Thelotremataceae (analysis 1) 

Our understanding of the circumscription of families in lichen-forming fungi is rather 

limited. This is due to lack of data, especially in crustose, tropical groups, but also due to 

persistence of traditional concepts based on oversimplifications of observed morphological 

diversity (Lumbsch, 2000, 2007). Traditionally, the distinction of families in lichenized fungi 

has been very coarse and based on simple, easy to recognize characters or correlations of few 

characters. Consequently, molecular data identified many traditionally circumscribed families 

as poly- or paraphyletic (e.g., Andersen & Ekman, 2004; Buschbom & Mueller, 2004; 

Ekman, 2001; Ekman & Jørgensen, 2002; Ekman & Wedin, 2000; Helms & al., 2003; 

Lumbsch & al., 2004; Mattsson & Wedin, 1999; Reeb & al., 2004; Schmitt & al., 2006; 

Wedin & al., 2000, 2002, 2005). However, it should be noted that some of the families 

circumscribed using morphological data, such as Parmeliaceae were supported as 

monophyletic by molecular data (e.g., Crespo & al., 2007). 

Although many questions remain open, it could be revealed by the combined analyses of 

two datasets employing nuclear and mitochondrial rDNA that Graphidaceae and 

Thelotremataceae in their present circumscription are both non-monophyletic. The distinction 

of both families was based on an overestimation of the importance of ascoma shape and 

development. The shape of the ascomata is easily observed and has therefore been used in 

traditional taxonomy (see also part 2. 1.). While most Graphidaceae have lirelliform 

ascomata, in some taxa these are rounded to ellipsoid, such as Acanthothecis hololeucoides, 

Diorygma confluens, Glyphis scyphulifera, Graphis muscicola, Leiorreuma hypomelaenum, 

and Phaeographis lobata (Staiger, 2002; Kalb & al., 2004). Lobate to almost lirelliform 

ascomata are known from some species of Chapsa (e.g., C. dilatata), Redingeria (e.g., R. 

glyphica), and Thelotrema (Frisch, 2006; Frisch & Kalb, 2006). Hence, the distinction of both 

families appears difficult in some cases, and the presence of these intermediate forms already 

made the use of this character set doubtful. Although these similarities are usually interpreted 

as homologies indicating close phylogenetic relationship, an alternative interpretation, i.e. that 

these similarities are due to parallelism, was expressed by Henssen (1976) and Henssen and 

Jahns (1973). These authors classified the two families in different suborders Ostropineae 

(Thelotremataceae) and Graphidineae (Graphidaceae) based on the differences in ascoma 

morphology and ascoma development. However, as examples for ascoma development in 

Graphidaceae they used Graphina mendax (=Diorygma junghuhnii) and Gyrostomum 

scyphuliferum (=Glyphis scyphulifera), two rather atypical examples since both genera 

produce partially anastomosing paraphyses in contrast to the mostly simple paraphyses of the 

other genera of Graphidaceae. Henssen & Jahns (ibid.) interpreted those as paraphysoids and 

compared the development to that of the unrelated Roccellaceae, but as shown by Staiger 

(2002) and Kalb & al. (2004), the different hamathecium anatomy in these genera is part of 

the anatomical and ontogenetic variation found in the family, which does not differ 

significantly from the variation found in Thelotremataceae (Frisch, 2006). 

Previous molecular studies already raised uncertainties in the distinction of the families. 

Grube & al., (2004) and Lumbsch & al., (2004a) found species of Thelotremataceae (the latter 

including Nadvornikia) being monophyletic, but nested within a paraphyletic Graphidaceae.


Frisch & al., (2006) found Graphidaceae and Thelotremataceae being polyphyletic, but the 

placement of Graphis spp. in a Thelotremataceae lineage lacked strong support. In a broad 

sampling of Graphidaceae including a few sequences of Thelotremataceae Staiger & al. 

(2006) found taxa of Thelotremataceae in three separate clades within Graphidaceae. 

However, the relationships of these clades were either unresolved or the relationships lacked 

significant support. Graphidaceae was not monophyletic with taxa of Dyplolabia and the 

paraphyletic Fissurina forming a separate well-supported monophyletic group with an 

unresolved relationship to the rest of Graphidaceae+Thelotremataceae and the outgroup. In an 

analysis addressing phylogeny of Lecanoromycetes using five gene regions, Miadlikowska & 

al. (2006) revealed a monophyletic Thelotremataceae (six species of two genera included) 

nested in a paraphyletic Graphidaceae. 

Here, it could be shown by the analysis of a dataset including over 50 Thelotremataceae 

and over 30 Graphidaceae species that the rejection of monophyly of the two families is 

legitimate. As a nomenclatural consequence of this analysis, it is suggested to reduce 

Thelotremataceae into synonymy with Graphidaceae, which will be formally proposed 

elsewhere. 

3. 4. 2. Phylogenetic relationship of genera within Graphidaceae (analysis 1, 2 and 3) 

The difficulties and controversial opinions in the distinction of major clades within 

thelotremataceaen Graphidaceae into genera were already discussed in part 2.1. As far as the 

Graphidaceae s. str. are concerned, a very similar discussion has evolved since Müller 

Argoviensis (1880a, 1882b) erected - prior to his introduction of a generic concept for 

Thelotremataceae (Müller Argoviensis, 1887) - the same ascospore-based classification for 

this group. The species were also distinguished in four major genera: brown and transversely 

septate ascospores: Phaeographis, brown and muriform ascospores: Phaeographina, hyaline 

and transversally septate ascospores: Graphis, hyaline and muriform ascospores: Graphina. 

However, this classification likewise persisted until recent times, although it was considered 

artificial by many workers (e. g. Archer, 2000; Santesson, 1952; Wirth & Hale, 1978). 

Several attempts towards a more natural classification in Graphidaceae, viz. the inclusion of 

ascomata-based characters and employment of molecular data, were presented in recent years 

(e. g. Kalb & Staiger, 2000; Kalb & al., 2004; Staiger, 2002; Staiger & al., 2006). As in the 

thelotrematoid taxa, numerous genera were newly introduced or resurrected, however, the 

classification is similarly far from being satisfactorily understood. 

Thus, in the combined gene analysis (analysis 1 – which is the only here presented analysis 

that includes this group), no further systematic conclusions can be drawn, except that the 

results of Staiger & al. (2006) could be confirmed, whereas Fissurina and Dyplolabia show a 

basal position within the Graphidaceae/Thelotremataceae clade, but again, this lacks support. 

The two genera were previously subsumed within Graphis s. lat. and Graphina, and 

reinstated by Kalb & Staiger (2001) and Staiger & Kalb (2000). Although the two genera look 

morphologically different, they have predominantly 3-septate or small muriform, I-negative 

ascospores as common characters. With the dataset at hand, the issue of generic concepts 

within Graphidaceae and Thelotremataceae cannot be addressed beyond the fact, that the 

current, recently introduced concepts (Frisch & al. 2006; Staiger 2002; Kalb & al. 2004) need 

further revision. One problem of this study is that several genera are not represented 

(Anomalographis, Anomomorpha, Gyrotrema, Ingvariella, Melanotrema, Phaeographopsis, 

Platythecium, Pseudoramonia, Redingeria, Reimnitzia, Sarcographina s. str., Thecaria), and 

of the 24 genera included, twelve are only represented by a single species. Also, type species 

are only included for 11 genera. Clearly, larger taxon sampling is needed to resolve the 

delimitation of most genera. Despite the shortcomings in taxon sampling, however, a number 

of genera were shown to be poly- or paraphyletic as currently circumscribed. This includes


the graphidaceous Graphis and Sarcographa and the thelotremataceous genera Chapsa, 

Myriotrema, Ocellularia, Thelotrema and Topeliopsis. 

In thelotremataceaen Graphidaceae, the only genus confirmed as monophyletic in two 

analyses (1 and 2), with more than one species represented, is Diploschistes. Although 

Thelotrema is strongly supported as monophyletic in analysis 3, with the extension of taxa 

beyond the Thelotrema s. str. group in the analyses 1 and 2, the genus as currently 

circumscribed happens to be polyphyletic. In the study of Frisch & al. (2006) Chapsa was 

monophyletic and sister to Leucodecton, both genera well supported. The placement of 

Leucodecton nested within Chapsa in analysis 1 may be due to poor taxon sampling, since we 

only included one species of Leucodecton, while the single-gene analysis of Frisch & al. 

(2006) included three taxa. In analysis 3, where six species of Chapsa are included as an 

outgroup, this genus appears well separated from Thelotrema but, firstly no other genus than 

Chapsa and Thelotrema is included, and, secondly, it does not form a monophyletic group. 

Neither Topeliopsis sensu Kantvilas and Vezda (2000) nor sensu Frisch & Kalb (2005) are 

monophyletic. However, Topeliopsis sensu Kalb (2001) is monophyletic in analysis 2 (also in 

analysis 1, but without support). The generic placement of Topeliopsis meridensis needs reevaluation. 

In both analyses it does not cluster with the type species T. muscigena or the 

strongly supported Topeliopsis s. str. group of analysis 2 respectively, but within Chapsa (and 

Thelotrema in analysis 2). Morphologically the species resembles other Chapsa species, but it 

differs in anatomical details, such as the sessile, rather urceolate apothecia, and a thick, brown 

proper exciple. However, the placement within Chapsa lacks support in both analyses and the 

generic placement of T. meridensis needs to be addressed with a wider sampling of Chapsa 

species. At the moment it is refrained from drawing any taxonomical conclusions. The former 

Topeliopsis species Melanotopelia toensbergii that is included in analysis 2 does also not 

cluster within the Topeliopsis s. str. group. When describing the new genus, Kantvilas and 

Vezda (2000) already mentioned that it was heterogeneous and that two species, viz. T. 

rugosa Kantvilas & Vezda and T. toensbergii Vezda & Kantvilas “may well be better placed 

in a separate genus altogether” (Kantvilas & Vezda 2000: 348). These two species were said 

to differ from the type species in having, at least in part, dark pigmented layers in the proper 

exciple (vs. hyaline proper exciple), a reddish reaction of the ascospores with iodine (vs. 

amyloid ascospores), and the presence of depsidones (vs. secondary metabolites lacking). 

Consequently Kalb (2001) restudied the generic circumscription of Topeliopsis and found that 

Topeliopsis s. str. differed in ascospores, being thin-walled (and non-amyloid, i.e. not reacting 

blue in iodine) in T. rugosa and T. toensbergii and thick-walled and amyloid in Topeliopsis 

s.str. He suggested to exclude the two deviating species from Topeliopsis and suggested they 

“probably belong to another undescribed genus” (Kalb 2001: 325), but did not suggested any 

nomenclatural consequences. Given the molecular support of analysis 2, the isolated position 

of these two species and the unique combination of morphological characters within 

thelotrematoid taxa, the new genus Melanotopelia is introduced here (see part 2. 9. 6., the 

formal description of the genus will be done elsewhere). 

Myriotrema in the current circumscription was also polyphyletic in the study by Frisch & 

al. (2006) and the results of analysis 1 confirm that its circumscription is in need of revision 

and additional data are necessary to evaluate its distinction from Ocellularia. The latter genus 

is currently poorly understood, our results confirm previous results that found this genus to be 

polyphyletic (Frisch & al., 2006, Lumbsch & al. 2004). Stegobolus has already been shown 

by Frisch & al. (2006) to be polyphyletic. 

In Thelotrema, a core group of Thelotrema s. str. can be recognized in all of the three 

analyses. This group largely agrees with the T. lepadinum group of Salisbury (1972) and 

Thelotrema subgen. Thelotrema of Matsumoto (2000). It is characterized by ascomata of the 

thelotremoid s. str. type as described in part 2. 5. 2. In analysis 1 it forms a sister group to T. 

crespoae and T. gallowayanum (within the clade P), in analysis 2 it is sister to T. weberi and


T. porinaceum in the Thelotrema sub-group of clade D, in analysis 3 it is resembled by the 

taxa of the clades B to G. The above mentioned taxa T. crespoae, T. gallowayanum, T. weberi 

and T. porinaceum (all but T. weberi included in the basal clade A of analysis 3), differ from 

the Thelotrema s. str. group by a perithecioid-thelotremoid type ascomata morphology (as 

described in part 2. 5. 2.). In addition, most of this species have large, muriform ascospores 

and either lack secondary metabolites or contain the norstictic acid chemosyndrome. 

However, these characters do also apply for T. saxatile and – to some extend – to T. 

monosporum, two species that cluster within the Thelotrema s. str. clades of analyses 1 and 3. 

In the analyses 1 and 2, five Thelotrema species cluster elsewhere (T. bicinctulum T. 

glaucopallens, T. myriocarpum, T. rugatulum, T. zebrinum). Some of those have 

morphological characteristics deviating from Thelotrema. The myriotremoid Thelotrema 

glaucopallens was already segregated as T. glaucopallens-group by Frisch (2006), without 

formally describing a separate genus, based on a fused to incompletely free exciple and the 

lack of lateral paraphyses. “Thelotrema” zebrinum is a member of the Leptotrema 

schizoloma-group, which is a rather well defined group of taxa that differ from the other 

members of Thelotrema by regenerating, distinctly layered ascomata and a carbonized exciple 

(see also part 2. 9. 13.). This group probably forms an independent genus, but further studies 

are needed to evaluate its systematic position. However, T. bicinctulum, T. myriocarpum and 

T. rugatulum fit morphologically into Thelotrema and their phylogenetic placement requires 

additional studies. 

In analysis 3, within the six well supported clades (clades B-G) of Thelotrema s. str., the 

two taxa of clade B (T. capetribulense and T. porinoides) agree in having emergent ascomata 

with a free proper exciple, transversely septate, hyaline, thick-walled, amyloid ascospores, 

and both contain the stictic acid chemosyndrome. The species of the clades C, D and F, T. 

diplotrema, T. nureliyum and T. subtile belong to a group of morphologically similar species, 

but also three species of clade E, viz. T. defossum, T. pseudosubtile and T. suecicum belong to 

this difficult to separate group. They share a similar thallus morphology, lack secondary 

metabolites and have hyaline, thick-walled, transversally septate ascospores. The other two 

taxa of clade E, T. lepadinum and T. rugatulum differ in having muriform ascospores. The 

distinction of T. subtile and T. suecicum as discussed by Purvis & al. (1995), which were 

treated as synonyms (Salisbury, 1972), is supported in our phylogenetic study. Also supported 

by the molecular results is the distinction of the new species T. pseudosubtile, which differs 

from T. subtile merely by subtle morphological differences and a more tropical distribution 

(see also under these species). It is assumed, that beyond the Australian collections from 

which it is currently only known, specimens from other tropical locations exist that were 

identified as T. subtile. Further, clade G includes four species that lack secondary metabolites 

and have brown – at least when mature – ascospores. One species has transversally septate 

ascospores, while eumuriform species are more common in this clade. This group includes 

some species, such as T. monosporum and T. saxatile, which are morphologically similar and 

due to different concepts of their circumscription, there was considerable confusion about the 

application of these names (Frisch & al. 2006).

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5. Appendices 322 

5. Appendices 

5. 1. Tentatively identified species 

Thelotrema cf. floridense Harris 

Some Florida Lichens: 98 (1990). Type: USA, Florida, Liberty County, Harris 1480 (US-holotype). 

Although it appears to be a characteristic species with a whitish, ecorticate thallus, 

regenerating ascomata with incurved, free exciple that is conspicuously raised above the 

hymenium, small, hyaline transversely septate ascospores and the psoromic acid 

chemosyndrome, the Hale collection from northern Queensland (see below) could not be 

identified with certainty since the type material was not available for study. So far T. 

floridense has not been reported from outside Florida. The Australian specimen agrees well 

with the description, however, it differs in ±split thalline rim margins. The identification is 

therefore tentative. 

SPECIMENS EXAMINED – Australia, Queensland: Near end of Black Mnt. Rd., 33 km WNW of Kuranda, Hale 

831909 (US). 

5. 2. Excluded or dubious names 

Ascidium attenuatum C. Knight 

Trans. New Zealand Inst. 15: 354 (1883). Phaeotrema attenuatum (Knight) Müll.Arg., Bull. Herb. Boissier, 

2: 74 (1894). Thelotrema attenuatum (Knight) Hellb., Bihang till Kgl. Svensk. Vetensk.-Akad. Handl. vol. 21, 

3(13): 78 (1896). Type: New Zealand, sine loco, C. Knight s.n. (WELT!-lectotype, selected by Galloway [1985: 

575] – without specimen, only drawing present). 

This name was considered synonymous to T. monosporum (Galloway, 1985). However, 

the type does not include a specimen but only a drawing of a single ascospore (selected as the 

type by Galloway) is available. Hence, A. attenuatum is here regarded as a nomen dubium. 

Chroodiscus mirificus (Kremp.) R.Sant. 

The report of this taxon from Australia (Lumbsch & Vezda, 1990) was based on 

erroneously identified collections of C. parvisporus. Chroodiscus mirificus is thus far not 

known from Australia. 

Conotremopsis weberiana Vezda 

Folia Geobot. Phytotax. 12: 314 (1977). 

This genus, which was listed as Thelotremataceae in McCarthy (2007) is currently placed 

in Stictidaceae (Eriksson, 2006).


Leptotrema lepadodes var. endochrysoides (Jatta) Zahlbr. 

Cat. Lich. Univ. 2: 636 (1923). Thelotrema lepadodes var. endochrysoides Jatta, Boll. Soc. Bot. Ital. 1910: 

256 (1911). Type: Australia, Tasmania, “Ad trunctum Pomaderris in monte Wellington, alt. 500 p.” (?NAP). 

The type of this variation was not available for study, although the herbarium in NAP was 

contacted. 

Leptotrema nitidulum Müll.Arg. 

Bull. Herb. Boissier 3: 315 (1895). Type: Australia, Queensland, 1887, Knight 22 (G!-holotype). 

This is a hitherto unpublished synonym of Stegobolus confluens (Kremp.) Mangold comb. 

nov. ined. 

Myriotrema microphthalmum (Müll.Arg.) Nagarkar & Hale 

Mycotaxon 35: 440 (1989). Bas.: Thelotrema microphthalmum Müll.Arg., Bull. Herb. Boissier 3: 314 

(1895). Type: Australia, Queensland, “Brisbane Shrubs”, Bailey 773 (G!-holotype; BRI-‘Shirley Book’, [p. 22, 

n. 18]!-isotype). 

This is a hitherto unpublished synonym of Ocellularia thelotremoides (Leight.) Zahlbr. 

Myriotrema minutulum (Hale) Hale 

Mycotaxon 11: 134 (1980). Bas.: Ocellularia minutula Hale, Smithson. Contr. Bot. 38: 24 (1978). Type: 

Panama, Canal Zone, Hale 43341 (US!-holotype). 

This taxon is not accepted as Myriotrema (=Ocellularia minutula Hale, acc. to Mangold & 

al. ined.). 

Thelotrema australiense Müll.Arg. 

Flora 70: 61 (1887). Type: Australia, New South Wales, Richmond River, Hodgkinson s.n. (M!-lectotype, 

selected by Hale [1970 in herb.]). 

This is a hitherto unpublished synonym of Ocellularia thelotremoides (Leight.) Zahlbr. 

Myriotrema microporellum (Nyl.) Hale 

The report of this taxon (McCarthy, 2007, as ‘Australian records’), was based on 

erroneously identified collections of Ocellularia thelotremoides. Myriotrema microporellum 

is thus far not known from Australia.


Thelotrema depressum Mont. {=Ocelularia depressa (Mont.) Hale, [non O. depressa 

Müll.Arg. =O. cavata (Ach.) Müll.Arg., (syn. nov.)]} 

The collection, on which the report of this taxon (Shirley, 1889b) is based on (Australia, 

Queensland, "On bark, 3-mile Shrub"), was not available. According to Shirley’s description 

(ibid.), however, the collection differs from the type of this species. 

Thelotrema expansum C.Knight [Phaeotrema expansum (C.Knight) Shirley] 

This is a hitherto unpublished synonym of Phaeographis lobata (Eschw.) Müll.Arg. 

Thelotrema inturgescens Müll.Arg. [in McCarthy (2007) as “T. intergescens”] 

This taxon is transferred to Ocellularia [=Ocellularia inturgescens (Müll.Arg.) Mangold 

comb. nov. ined.]. 

Thelotrema puniceum (Müll.Arg.) Makhija & Patw. [Chapsa punicea (Müll.Arg.) Caceres & 

Lücking] 

This taxon is transferred to Ocellularia [=Ocellularia punicea (Müll.Arg.) Mangold comb. 

nov. ined.]. 

Thelotrema rimulosum Müll.Arg. 

This taxon is transferred to Hemithecium [=Hemithecium rimulosa (Müll. Arg.) Mangold, 

Lumbsch & Kalb comb. nov. ined.]. 

Thelotrema subgranulosum Jatta 

Boll. Soc. Bot. Ital. 1910: 256 (1911). Type: Australia, Tasmania, Mt. Wellington, s. col. [NAP- no reply 

from herbarium]. 

The type of this taxon was not available for study. 

Thelotrema trypethelioides C.Knight ex F.M.Bailey [Phaeotrema trypethelioides (C.Knight 

ex F.M.Bailey) Shirley]. 

This is a hitherto unpublished synonym of Stegobolus fissus (Nyl.) A.Frisch. 

Tremotylium australiense Müll.Arg. / Tremotylium nitidulum Müll.Arg. 

Material of these species was not examined. The genus Tremotylium was transferred to 

Minksia (Roccellaceae) by Makhija & Patwardhan (1995) based on the type species 

Tremotylium angolense (=Minksia angolensis).


5. 3. List of synonyms 

The following is a list of all accepted synonyms of the present treatment, alphabetically 

arranged after the genus name of the basionym: 

Anthracothecium oligosporum Müll.Arg. =Leucodecton compunctellum 

Ascidium manosporum Knight =Thelotrema saxatile 

A. octolocularis Knight =Thelotrema bicinctulum 

Endocarpon baileyi Stirt. =Leptotrema wightii 

Graphis phlyctidea Vain. =Chapsa leprieurii 

G. subnivescens Nyl. =Chapsa leprieurii 

Leptotrema aemulum Müll. Arg. =Thelotrema lepadodes 

L. albocoronata Knight =Thelotrema leucophthalmum 

L. bisporum Szatala =Thelotrema lepadodes 

L. compunctum var. purpuratum Müll. Arg. =Leucodecton occultum 

L. deceptum Hale =Leucodecton compunctellum 

L. diffractum Müll. Arg. =Leucodecton subcompunctum 

L. inclusum Zahlbr. =Leucodecton subcompunctum 

L. mastoideum Müll.Arg. =Reimnitzia santensis 

L. nitidulum Müll.Arg. =Stegobolus confluens 

L. polycarpum Müll.Arg. =Leucodecton subcompunctum 

L. polyporum Riddle =Myriotrema phaeosporum 

Leucodecton biokense A. Frisch =Leucodecton compunctellum 

Myriotrema decorticatum Hale =Leucodecton subcompunctum 

M. multicavum Hale =Myriotrema rugiferum 

M. nuwarense Hale =Leucodecton compunctellum 

M. subcostaricense Sipman =Myriotrema glaucophaenum 

Ocellularia alba var. caesiascens Räs. =Chapsa astroidea 

O. annulosa Müll.Arg. =Thelotrema lacteum 

O. bonplandiae var. obliterata Müll. Arg. =Thelotrema suecicum 

O. costaricensis Müll. Arg. =Myriotrema glaucophaenum 

O. cricota F.Wilson =Thelotrema lacteum 

O. demersa Müll.Arg. 

[nom. nov. pro Pyrenula clandestina Fée] =Thelotrema bicinctulum 

O. galactina Zahlbr. =Myriotrema microporum 

O. jugalis Müll.Arg. =Thelotrema subtile 

O. platychlamys Müll. Arg. =Thelotrema porinoides 

O. turgidula Müll. Arg. =Thelotrema diplotrema 

O. zeorina Müll.Arg. =Thelotrema lacteum 

Phaeotrema apertum C. W. Dodge =Chapsa platycarpa 

P. consimile Müll.Arg. =Thelotrema lacteum 

Thelotrema aemulans Kremp. =Thelotrema lepadinum


T. albescens Vain. =Chapsa indica 

T. albidiforme Leight. =Thelotrema porinoides 

T. argenteum Müll.Arg. =Fibrillithecis halei 

T. australiense Müll.Arg. =Ocellularia thelotremoides 

T. clandestinum f. remanens Nyl. =Myriotrema clandestinum 

T. colobicum Nyl. =Chapsa leprocarpa 

T. compunctum f. portoricensis Vain. =Leucodecton occultum 

T. compunctum var. antillarum Vain. =Leucodecton occultum 

T. compunctum var. praiense Vain. =Leucodecton occultum 

T. confluens Vain. =Chapsa leprieurii 

T. crassulum Nyl. =Myriotrema microporum 

T. diminitum Hale =Fibrillithecis halei 

T. disciforme Leight. =Thelotrema lepadodes 

T. dissultum Hale =Thelotrema cupulare 

T. elachistoteron Leight. =Leucodecton compunctellum 

T. emergens Vain. =Myriotrema glaucophaenum 

T. exalbidum Stirt. =Thelotrema pachysporum 

T. exanthismocarpum Leight. =Thelotrema porinoides 

T. expansum C. Knight =Phaeographis lobata 

T. flavescens Darb. =Thelotrema lepadinum 

T. galactinum Vain. =Thelotrema pachysporum 

T. gibbosum H. Magn. =Fibrillithecis halei 

T. heterosporum C. Knight =?Reimnitzia santensis (see under this species 

for details) 

T. homothecium Vain. =Thelotrema porinoides 

T. hypomelaenum Müll. Arg. =“Leptotrema” schizoloma 

T. indicum Hale =Topeliopsis muscigena 

T. irosinum Vain. =Myriotrema desquamans 

T. laevius Vain. =Myriotrema trypaneoides 

T. leucastrum Tuck. =Chapsa leprieurii 

T. leucastrum var. difforme Tuck. =Chapsa leprieurii 

T. leucohymenium Zahlbr. =Myriotrema viridialbum 

T. leucophthalmum var. lacerata Räs. =Chapsa megalophthalma 

T. limae Vain. =Thelotrema pachysporum 

T. microglaenoides Vain. =Leucodecton compunctellum 

T. microphthalmum Müll.Arg. =Ocellularia thelotremoides 

T. monospermum Harris =Thelotrema saxatile 

T. monosporoides Nyl. =Thelotrema saxatile 

T. monosporum var. patulum Nyl. =Thelotrema lepadodes 

T. monosporum var. subgemium Nyl. =Leucodecton compunctellum 

T. myrioporoides Müll. Arg. =Myriotrema viridialbum 

T. obconicum Raes. =Thelotrema lepadinum 

T. obovatum Stirt. =Thelotrema porinoides 

T. pachystomum ssp. piluliferum Tuck. =Fibrillithecis halei 

T. palmense Vain. =Thelotrema pachysporum 

T. platycarpellum Vain. =Chapsa astroidea 

T. platycarpoides Tuck. =Chapsa platycarpa 

T. platysporum Harm. =Fibrillithecis halei 

T. pycnophragmium Nyl. =Chapsa indica 

T. ravenelii Tuck. =Leptotrema wightii 

T. reclusum Kremp. =Leucodecton compunctellum


T. rimulosum Müll.Arg. =Hemithecium rimulosa 

T. sitianum Vain. =Thelotrema lacteum 

T. steyermarkii Hale =Myriotrema viridialbum 

T. subcaesium Nyl. =Myriotrema rugiferum 

T. subconcretum Leight. =Leptotrema wightii 

T. subcrassulum Vain. =Myriotrema olivaceum 

T. subterebrans Nyl. =Myriotrema trypaneoides 

T. terebrans Nyl. =Thelotrema bicinctulum 

T. terebratulum Nyl. =Myriotrema clandestinum 

T. trypethelioides C.Knight =Stegobolus fissus 

T. vernicosum Zahlbr. =Fibrillithecis halei 

Topeliopsis corticola Kalb =Topeliopsis decorticans 

T. muscicola Kantv. & Vezda =Topeliopsis muscigena 

T. vezdae Kalb =Topeliopsis subdenticulata 

Tylophoron diplotylium Nyl. =Nadvornikia hawaiensis 

Urceolaria compuncta Ach. =Leucodecton occultum

LEBENSLAUF 

Persönliche Daten 

Name: Armin Mangold 

Geburtsdatum: 17.04.1972 / Stuttgart 

Anschrift: Hasenheide 56, 10967 Berlin 

Tel.: 030/22808214 

eMail: arminmangold@gmail.com 

Familienstand: verheiratet 

Staatsangehörigkeit: deutsch 

Schulbildung 

1988 - 1991 Louis Leitz Schule Stuttgart (vorm. 

„Wirtschaftsgymnasium Stuttgart-Feuerbach“) 

Abschluss: Abitur 

Zivildienst und universitäre Ausbildung 

10/1991 - 04/1993 Zivildienst 

10/1993 - 10/1996 Studium Diplom-Biologie an der RWTH Aachen 

Abschluss: Vordiplom 

10/1996 - 01/2003 Studium Lehramt Biologie und Kunst 

(Sekundarstufe I und II) an der Universität Duisburg- 

Essen 

Abschluss: 1. Staatsexamen 

Seit 01/2003 Promotion an der Universität Duisburg-Essen. 

Größtenteils durchgeführt am Field Museum Chicago 

(USA), betreut von Dr. T. H. Lumbsch. 

Berlin, 26.01.2008

Erklärung: 

Hiermit erkläre ich, gem. § 6 Abs. 2, Nr. 6 der Promotionsordnung der Math.-Nat.- 

Fachbereiche zur Erlangung des Dr. rer. nat., dass ich die vorliegende Dissertation 

selbständig verfasst und mich keiner anderen als der angegebenen Hilfsmittel 

bedient habe. 

Essen, den _________________ ______________________________ 

Erklärung: 


Fachbereiche zur Erlangung des Dr. rer. nat., dass ich das Arbeitsgebiet, dem das 

Thema “Taxonomic studies on members of thelotrematoid Ostropales (lichenized 

Ascomycota) in Australia” zuzuordnen ist, in Forschung und Lehre vertrete und den 

Antrag von Armin Mangold befürworte. 

Essen, den _________________ ____________________________________ 

Erklärung: 


Fachbereiche zur Erlangung des Dr. rer. nat., dass ich keine anderen Promotionen 

bzw. Promotionsversuche in der Vergangenheit durchgeführt habe und dass diese 

Arbeit von keiner andern Fakultät abgelehnt worden ist. 

Essen, den _________________ _________________________

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