Four new -associated Trimmatostroma species (hyphomycetes)

Paul Diederich1, Uwe Braun2, Bettina Heuchert2 & Damien Ertz3

1 Musée national d’histoire naturelle, 25 rue Munster, L-2160 Luxembourg, Luxembourg ([email protected]) 2 Martin Luther University, Institute of Biology, Department of Geobotany and Botanical Garden, Herbarium, Neuwerk 21, D-06099 Halle (Saale), Germany ([email protected] and bettina.heuchert@botanik. uni-halle.de) 3 Jardin Botanique National de Belgique, Domaine de Bouchout, B-1860 Meise, Belgique ([email protected])

Diederich P., U. Braun, B. Heuchert & D. Ertz, 2010. Four new lichen-associated Trimmato­ stroma species (hyphomycetes). Bulletin de la Société des naturalistes luxembourgeois 111: 47-55.

Abstract. Four new lichen-associated species of Trimmatostroma s. lat. are described. T. dendrographae is found on Dendrographa in California (U.S.A.), T. hierrense on cf. Arthonia endlicheri in El Hierro (Canary Islands), T. lecanoricola on saxicolous Lecanora in Peru; T. quercicola develops over bark of Quercus and is facultatively lichenicolous on degenerate lichen thalli in Belgium and Luxembourg. A key to the known lichenicolous species of Trimmatostroma is provided.

Key words. Anamorphic fungi, Intralichen, lichenicolous, Taeniolella.

1. Introduction Helotiales (Crous et al. 2007). Other spe- cies were transferred to the new genus Cat­ The genus Trimmatostroma Corda (1837), enulostroma Crous & U. Braun (Crous et with T. salicis Corda as type species, was al. 2007). They are morphologically close introduced for bark-inhabiting saprobic to Trimmatostroma s. str., but belong to the hyphomycetes. This genus is character- Teratosphaeriaceae (Capnodiales). How- ized by having simple, little differenti- ever, even Catenulostroma is in its cur- ated conidiophores and conidiogenous rent circumscription probably not mono- cells as well as polymorphous conidia phyletic (Crous et al. 2009). formed in irregular basipetal chains (Ellis During the course of monographic stud- 1971, 1976). T. lichenicola M.S. Christ. & ies on lichenicolous Taeniolella S. Hughes D. Hawksw. (in Hawksworth 1979) was species, several collections provisionally the first lichenicolous species assigned to named “Taeniolella sp.” were encountered this genus. Hawksworth & Cole (2002) that turned out to be Trimmatostroma-like. revised and reassessed T. lichenicola and Species of Taeniolella are easily distinguish- similar lichenicolous hyphomycetes and able by having superficial, semimacrone- introduced the new genus Intralichen D. matous conidiophores, conidia formed Hawksw. & M.S. Cole, mainly differentiated in acropetal chains, and lacking multicel- from Trimmatostroma by its lichenicolous lular aggregations of conidial cells, which habit, an immersed mycelium, entirely are common in Trimmatostroma. As no immersed micronematous conidiophores, sequence data are available for these spe- and pale, smooth-walled conidia with few cies and as conidiomata are not stromatic septa. Based on molecular sequence anal- (the type species T. salicis and related spe- yses, Trimmatostroma proved to be very cies have sporodochial conidiomata with heterogeneous and must probably be con- a basal stroma), an inclusion in Trimma­ fined to T. salicis and a few closely allied tostroma might be questionable. On the species phylogenetically pertaining to the other hand, the formation of stromata

Bull. Soc. Nat. luxemb. 111 (2010) 47 and conidiophores in sporodochia is often colous taxa tentatively in Trimmatostroma unreliable as distinctive generic charac- s. lat. The final generic position of licheni- ter in hyphomycete genera. However, no colous Trimmatostroma-like hyphomycetes other suitable genus is available. There- depends on the phylogenetic affinity of the fore, we prefer to include the new licheni- taxa concerned.

Fig. 1. Trimmatostroma dendrographae. A–C, Infected thalli of Dendrographa. D, Conidiophore with chains of conidia, in water. E, Same, in KOH. F–J, Chains of conidia, in water. K, Conidiophore, in water. A: California, Monterey, Crocker Grove; B–C, F–K: holotype; D–E: Diederich 16767. Scale bars: B = 1 mm, C = 0.5 mm, D–K (same scale bar) = 10 μm.

48 Bull. Soc. Nat. luxemb. 111 (2010) 2. Material and Methods subtruncate. Conidia in simple or rarely branched, irregular, rarely disarticulating, The specimens examined are deposited in B, basipetal chains, shape and size very vari- BR, HAL, NY, UCR and UPS, and in the pri- able, mostly ellipsoid-ovoid to subcylindri- vate herbarium of P. Diederich. For micro- cal or oblong, transversely 0–pluriseptate, scopical examination, small fragments were rarely dictyosporous, ends more or less examined in water, after light pressure on rounded, c. 7–25 × 5.5–8.5 µm, sometimes the cover glass, and in 5% KOH. forming irregular aggregations, medium to dark brown or olivaceous-brown, distinctly olivaceous in KOH, wall 0.8–1.2 μm thick, 3. Results wall rugose-rimulose. Trimmatostroma dendrographae Diederich, This species is distinguished from T. leca­ Ertz, U. Braun & Heuchert sp. nov. Fig. 1 noricola by olivaceous conidia in KOH, from T. hierrense by rugose-rimulose conidia and MycoBank 518957. from T. quercicola by narrower conidia that Trimmatostromatis quercicolae simile, sed cat- are not visible macroscopically. Attempts to enis conidiorum macroscopice non distinctis, culture freshly collected material failed. thallu hospitis plus minusve fuscescens, conidiis olivaceo-bruneis in aqua, intense olivaceis in Ecology, hosts and distribution: All known KOH, 5.5–8.5 µm latis, pariete usque ad 1.2 µm specimens are lichenicolous on epiphytic crasso. Differt a Trimmatostromate salicis et T. and saxicolous thalli of Dendrographa, betulino habitu lichenicola, stromatibus et spo- including crustose, sorediate thalli that are rodochiis nullis, coloniis dispersis supra thallos considered to be the primary morph of the Dendrographae, conidiis olivaceo-brunneis, oli- host, before the fruticose structures appear. vaceis in KOH, 0–pluriseptatis, 7–25 × 5.5–8.5 In many specimens, fruticose thalli start µm, pariete rugoso-rimuloso. growing, but are so heavily parasitized by Type: U.S.A., California, Monterey Co., Carmel, this hyphomycete that they just reach up to Point Lobos State Reserve, 30 m alt., on Den­ drographa sp., on , 18 Jul. 1 cm in length and then die. This species is 2008, P. Diederich 16789 & D. Ertz (BR – holo- known only from the coast of California, and type; HAL, NY, UCR, herb. Diederich – isotypes). it appears to be rather common in localities Colonies lichenicolous over the thallus of where the host Dendrographa is present. Dendrographa, usually covering the entire Additional specimen examined (all on Den­ surface of the host thallus, blackish, mac- drographa): USA, California, Monterey Co.: roscopically appearing as a finely granu- same locality as the type, on Cupressus macro­ carpa, 2008, Ertz 12482, 12487 & Diederich (BR); lar mixture of blackish conidia and white ibid., sheltered rock face near the sea, 2008, Ertz host hyphae, the whole appearing pale to 12441, 12453 & Diederich (BR); SW of Monterey, dark greyish at a low magnification (e.g., 17 Mile Drive, Crocker Grove, alt. 15 m, on C. in the field), individual chains of conidia macrocarpa, 2008, Diederich 16767 & Ertz 12462 hardly visible. Mycelium internal, hyphae (BR, herb. Diederich); along coast S of Asilomar, sparingly branched, 2.5–3.5 µm wide, sub- China Rock, alt. 5 m, on rock outcrop on the hyaline to pale brown, septate, thin-walled, beach, 2008, Ertz 12480 & Diederich (BR). rugose. Stroma lacking. Conidiophores usu- ally immersed, occasionally somewhat Trimmatostroma hierrense Diederich & erumpent, micronematous, c. 20–70 × 2.5–4 Ertz sp. nov. Fig. 2 µm, hyphal filaments gradually develop- MycoBank 518958. ing into fertile threads by becoming some- Differt ab omnibus speciebus Trimmatostromatis what wider and darker, with slightly thicker conidiis transverse septatis et parietibus conidi- wall, but differentiation between hyphae orum unilateraliter incrassatis et fuscatis. and conidiophores very difficult, rugose. Type: Spain, Canary Islands, El Hierro, E of Conidiogenous cells integrated, terminal, Frontera, trail between Frontera and ‘mirador de monoblastic (i.e., the terminal cell function- Jinama’ (part below ‘Fuente de Tincos’), 27°45’29” ing as conidiogenous cell), c. 5.5–9 × 3.5–4 N, 17°59’29” W, ‘fayal brezal’ forest and rocky µm, conidiogenous loci undifferentiated, outcrops, on a root under a rock along trail, on cf.

Bull. Soc. Nat. luxemb. 111 (2010) 49 Fig. 2. Trimmatostroma hierrense (holotype). A–B, Infected thalli of cf. Arthonia endlicheri. C, Conidia and conidial chains, in water. D–F, Same, in KOH (conidiophores visible in D). Scale bars: A = 1 mm, B = 0.2 mm, C–F = 10 μm.

Arthonia endlicheri, 31 Aug. 2009, D. Ertz 13946 and conidiophores very difficult, smooth. (BR – holotype; herb. Diederich – isotypes). Conidiogenous cells integrated, terminal, Colonies lichenicolous over the thallus of cf. monoblastic (i.e., the terminal cell func- Arthonia endlicheri, covering large parts of tioning as conidiogenous cell), c. 5.5–10 × the host thallus, blackish, macroscopically 3.5–4.5 µm, conidiogenous loci undifferenti- appearing as a finely granular mixture of ated, subtruncate. Conidia in simple or rarely blackish conidia and white host hyphae, the branched, irregular, often disarticulating, whole appearing dark greyish to blackish at basipetal chains, shape and size very vari- a low magnification (e.g., in the field), indi- able, mostly ellipsoid-ovoid to subcylindrical vidual chains of conidia hardly visible. Myc­ or oblong, often with irregular convex and elium internal, hyphae sparingly branched, concave regions, transversely 0–pauciseptate, 2–2.5 µm wide, subhyaline to pale brown, ends more or less rounded, c. 7.5–15 × 3.5– septate, thin-walled, smooth. Stroma lacking. 6.5 µm, pale to medium brown, partly with Conidiophores usually immersed, occasion- thickened walls and then dark brown, oliva- ally somewhat erumpent, micronematous, up ceous brown in KOH; wall c. 0.5 μm thick in to 40 μm long and 2.5–3.5 µm wide, hyphal pale areas, c. 1 μm in darker areas, smooth. filaments gradually developing into fertile Macroscopically, the new species looks very threads by becoming somewhat wider and similar to Trimmatostroma dendrographae, darker, but differentiation between hyphae entirely darkening large areas of the host

50 Bull. Soc. Nat. luxemb. 111 (2010) thallus. Microscopically, it is distinguished B. Ellis, another species with an unevenly by conidia frequently with a dark and thick thickened and pigmented conidial wall, outer wall limited to one side of the conid- but in that species conidia are multicellular ium (resembling conidia of Milospium with transverse and longitudinal septa (Ellis graphideorum). Furthermore, conidia are 1976). usually smoother than in T. dendrographae, Ecology, hosts and distribution: The new best seen in KOH. Because of the unevenly species covers large parts of the thallus of pigmented conidial wall, T. hierrense might a white crustose lichen with be mistaken for a species of Milospium. reacting C+ red, that probably belongs to However, conidia of the type species, M. Arthonia endlicheri, growing on the root of graphideorum (Nyl.) D. Hawksw., have a a tree along a trail. Infected thalli become very different shape, being irregularly lobed entirely dark greyish black. This species is and aseptate, and they are never catenate. known only from the type locality on the isle Conidia of M. deslooveri Diederich & Sérus. of El Hierro (Spain, Canary Islands). (Sérusiaux et al. 1999) and M. planorbis (Aptroot & Sipman 2001; holotype from B examined) are strongly curved, multi- Trimmatostroma lecanoricola Diederich, cellular, often with incomplete septa, and Etayo, U. Braun & Heuchert sp. nov. Fig. 3 never catenate; furthermore, both species MycoBank 518959. are lichenized, not lichenicolous. The new Trimmatostromatis dendrographae et T. quercico­ species could also be confused with Trim­ lae simile, sed catenis conidiorum disarticulan- matostroma scutellare (Berk. & Broome) M. tium aurantiaco-brunneis in KOH, nec umquam

Fig. 3. Trimmatostroma lecanoricola (holotype). A, Thallus of saxicolousLecanora with young infection. B, Thallus with older infection. C, Chains of conidia, in water. D, Same, in KOH. E, Disarticulating chain of conidia, in water. F, Conidium in surface view, showing verrucose, mosaic-like outer layer of conidial wall, in water. Scale bars: A–B = 0.5 mm, C–F = 10 μm.

Bull. Soc. Nat. luxemb. 111 (2010) 51 olivaceo-brunneis, et partibus atrioribus paginae Trimmatostroma quercicola Diederich, U. conidiorum minoribus in conidiis vetustioribus Braun & Heuchert sp. nov. Figs 4–5 rimulosis. MycoBank 518960. Type: Peru, Dep. La Libertad, Prov. Trujillo, Cerro Malabrigo, SW of Puerto Chicama, on saxicolous Trimmatostromatis dendrographae simile, sed Lecanora, 100–150 m alt., 9 Apr. 1981, R. Santes­ catenis conidiorum macroscopice expedite con- son & A. Tehler P128:1 (UPS – holotype). spicuis, irregulariter reticuloideis, conidiis badiis in aqua et olivaceo-brunneis in KOH, 5–15 µm Colonies lichenicolous, over the thallus and latis, pariete usque ad 2 μm crasso. Differt a T. apothecia of a saxicolous Lecanora; when lecanoricola catenis conidiorum raro disarticula- young, appearing as irregular, branched, tis, conidiis olivaceo-brunneis in KOH, et strato blackish filaments (representing conidial exteriore conidiorum obscuriore in conidiis chains), soon aggregating in very dense vetustioribus rimulosis. tufts of conidial chains, the infected parts of Type: Luxembourg, Vogelsmühle, valley of Haler- the host becoming entirely blackish. Myc­ baach, left side of river, on bark ofQuercus , possi- elium internal, hyphae sparingly branched, bly associated with sterile crustose , 9 Dec. 1.5–3 µm wide, subhyaline to pale brown, 2007, P. Diederich 16723 (BR – holotype; HAL, septate, thin-walled, smooth. Stroma lack- herb. Diederich – isotypes). ing. Conidiophores usually immersed, occa- Colonies over bark of Quercus, frequently sionally somewhat erumpent, micronema- overgrowing and probably parasitizing tous, c. 20–40 × 2–4 µm, hyphal filaments degenerate crustose lichen thalli or corti- gradually developing into fertile threads colous , effuse, loose to dense, fumose- by becoming somewhat wider and darker, black, macroscopically appearing as rela- with slightly thicker wall, but differentiation tively short, prostrate, irregularly formed between hyphae and conidiophores very and branched, often agglomerated conidial difficult. Conidiogenous cells integrated, ter- chains. Mycelium internal, hyphae sparingly minal, monoblastic (i.e., the terminal cell branched, 1–3.5 µm wide, subhyaline to pale functioning as conidiogenous cell), c. 4–7 brown, septate, thin-walled, smooth. Stroma × 3–4.5 µm, conidiogenous loci undiffer- lacking. Conidiophores usually immersed, entiated, subtruncate. Conidia in simple or occasionally somewhat erumpent, microne- rarely branched, irregular, easily disarticu- matous, c. 5–40 × 2–6 µm, hyphal filaments lating, basipetal chains, often subspherical gradually developing into fertile threads and 1-septate, 6.5–10 × 6–8 μm, or oblong, by becoming somewhat wider and darker, transversely pluriseptate to dictyosporous, with slightly thicker wall, but differentiation c. 11–24 × 5–9 µm, medium to dark brown, between hyphae and conidiophores very orange brown in KOH, wall 0.5–1.2 µm difficult. Conidiogenous cells integrated, thick, rugose-rimulose to irregularly verru- terminal, monoblastic (i.e., the terminal cose, ends more or less rounded. cell functioning as conidiogenous cell), c. This species is readily distinguished from 5–10 × 3–4 µm, conidiogenous loci undif- the other taxa treated here by conidial chains ferentiated, subtruncate. Conidia in simple becoming orange brown in KOH, never with or rarely branched, irregular, occasionally an olivaceous tinge, and by chains of disar- disarticulating, basipetal chains, shape and ticulating conidia. Furthermore, when the size very variable, globose or subglobose, outer layer of the conidial wall splits with 0–1(–2)-septate, 4–9 µm diam., or ellipsoid- age, the resulting darker patches are often ovoid, subcylindrical, oblong, 1–4(–5)-sep- smaller than in the other species, giving the tate, transversely septate to dictyosporous, c. conidial surface a mosaic-like appearance. 6–30 × 5–15 µm, sometimes forming irregu- Ecology, host and distribution: This spe- lar aggregations, medium to dark brown, cies is known only from the type collection olivaceous-brown in KOH, wall thick, up to made in Peru on an unidentified saxicolous 2 µm, occasionally distinctly two-layered, Lecanora. Infected host thalli and apothecia i.e., with a distinct, paler inner layer (but not are damaged, suggesting that the fungus is distoseptate), wall rugose-rimulose to irreg- pathogenic. ularly verrucose, ends more or less rounded.

52 Bull. Soc. Nat. luxemb. 111 (2010) Amongst the new species described here, T. Pirozynski 1965). Although the type of this quercicola is macroscopically rather distinct, species has never been restudied and the as conidial chains are easily visible at a strong species never been reported after its original magnification (× 40), appearing as an irregu- description, it seems to differ clearly from lar, branched net of relatively thick hyphae the new T. quercicola by the pulvinate, stro- overgrowing the substratum. Microscopi- matic conidiomata, the different ecology cally, the species has much thicker conidial (rotten wood, versus bark of Quercus) and chains than the three others. It is chemically the mostly larger conidia. distinct from T. lecanoricola by the K+ oliva- Ecology, hosts and distribution: The new ceous conidial chains and differs slightly from species is known from two collections on T. dendrographae in having brown conidia (in the bark of old Quercus. As conidiophores water) without a distinct olivaceous tinge. at least occasionally overgrow and possibly Trimmatostroma quercinum (Hoffm.) Höhn. parasitize reduced lichen thalli, the species (syn. Phragmotrichum quercinum Hoffm.) may be considered as facultatively licheni- was described from rotten manufactured colous. It is probably widespread, but over- wood of Quercus in Germany. The species looked in similar habitats, and currently forms black, superficial, dispersed, orbicu- known from Belgium and Luxembourg. lar, pulvinate stromata, and oblong, muri- Additional specimen examined: Belgium, SE of cate, transversally septate to irregularly Buzenol, on bark of Quercus, possibly associated muriform, indistinctly catenate conidia with sterile crustose lichens, 19 June 1984, P. Die­ 25–30 × 12–15 μm (Migula 1921, Sutton & derich 5576 (herb. Diederich).

Fig. 4. Trimmatostroma quercicola (holotype). A, Conidiophores with and without conidia. B, Plagiotropous conid- ial chain. C, Conidia. Scale bar = 10 µm. U. Braun del.

Bull. Soc. Nat. luxemb. 111 (2010) 53 Fig. 5. Trimmatostroma quercicola. A, Overgrowing degenerate lichen thalli on Quercus bark. B, Saprobic over Quer­ cus bark. C, Squash preparation of infected lichen thallus, in water. D, Chains of conidia, in water. E, Same, in KOH. A, C: holotype; B, D–E: Diederich 5576. Scale bars: A–B = 200 μm, C–E = 10 μm.

4. Discussion Conidia are superficially similar to those of Spilodochium Syd. (Ellis 1971, 1976). How- Hawksworth (1979) described the licheni- ever, the genus Spilodochium is easily dis- colous Trimmatostroma lichenicola M.S.Christ. tinguishable by its erumpent pseudoparen- & D. Hawksw., intrahymenial in Candelariella chymatic stromata, conidiophores arranged vitellina. Later, Hawksworth & Cole (2002) in sporodochia and conidia produced in introduced the new generic name Intrali­ branched acropetal chains. The entire habit chen for this and several similar species, and of the new species is rather Intralichen-like. no lichenicolous species was subsequently However, species of Intralichen have paler, accepted in Trimmatostroma. smaller, smooth-walled, mostly 0–2-celled The four new species share a large number conidia, and conidiophores are usually of characters. They all have dark, micro- entirely immersed in lichen thalli or apoth- nematous conidiophores not arranged in ecia, most commonly being intrahymenial, sporodochia, lack stromatic hyphal aggrega- with only the mature conidia arising at the tions, have integrated conidiogenous cells, surface (Hawksworth & Cole 2002). On the and conidia produced in simple or rarely other hand, the inclusion of these species branched basipetal chains, 1–pluriseptate, in Taeniolella is no alternative, since spe- irregular in size and form, with an often cies of this genus do not form dictyosporous rough surface, the outer layer of the wall conidia, and its conidial chains are acropetal. often splitting with age, resulting in pale Microscopically, the new species are similar to medium brown conidia covered by dark to Trimmatostroma salicis, the type species, brown remnants of the outer layer, or une- and T. betulinum (Corda) S.Hughes, from venly pigmented. Furthermore, they all obli- which they are distinguished by the absence gately or facultatively parasitize lichen thalli. of a basal stroma and lacking pulvinate spo-

54 Bull. Soc. Nat. luxemb. 111 (2010) rodochia (Crous et al. 2007). The latter two 3. Conidia with a dark outer wall splitting species have been shown to belong to the with age, but more evenly surrounding the Helotiales (Crous et al. 2007). Many Trim­ entire conidia, wall rugose; lichenicolous on matostroma-like fungi have recently been Dendrographa ...... T. dendrographae transferred to the new genus Catenulos­ troma in the Teratosphaeriaceae [Capnodi- ales] (Crous et al. 2007), which is, however, Acknowledgements no alternative either, since this genus is also We are much obliged to the curators of B and characterized by having stromatic conidi- UPS for the possibility to examine collections omata. The introduction of a new genus for from their herbaria, and to Prof. Walter Gams for the new lichenicolous species might be the critically commenting on the manuscript. best solution, but in the absence of molecu- lar data, i.e., without knowledge of the phy- References logenetic affinity, we prefer to assign the new Aptroot, A. & H. J. M. Sipman, 2001. New Hong species tentatively to Trimmatostroma s. lat. Kong lichens, ascomycetes and lichenicolous fungi. Journal of the Hattori Botanical Labora­ tory 91: 317–343. 5. Identification key for the known Corda, A. K. J., 1837. Icones fungorum hucusque lichenicolous species of Trimmato- cognitorum. Vol. 1. J. G. Calve, Praha. stroma Crous, P. W., U. Braun, J. Z. Groenewald, 2007. Mycosphaerella is polyphyletic. Studies in 1. Conidia orange brown in KOH; conidial Mycology 58: 1–32. chains frequently disarticulating; outer layer of conidial wall splitting with age and then some- Crous, P. W., C. L. Schoch, K. D. Hyde, A. R. times divided in many relatively small patches, Wood, C. Gueidan, G. S. de Hoog & J. Z. looking mosaic-like (Fig. 3F); lichenicolous on Groenewald, 2009. Phylogenetic lineages saxicolous Lecanora...... T. lecanoricola in the Capnodiales. Studies in Mycology 64: 17–47. 1. Conidia olivaceous-brown in KOH; Ellis, M. B., 1971. Dematiaceous hyphomycetes. conidial chains rarely disarticulating; outer Commonwealth Mycological Institute, Kew. layer of conidial wall either smooth (Fig. 2) or splitting with age and then divided into Ellis, M. B., 1976. More dematiaceous hyphomyc- etes. Commonwealth Mycological Institute, several larger patches (Figs 1 & 5D) ...... 2 Kew. 2. Conidial chains easily visible macroscop- Hawksworth, D. F., 1979. The lichenicolous ically (× 40), appearing as an irregular net of hyphomycetes. Bulletin of the British Museum blackish hyphae; conidia reddish brown in (Natural History), Botany 6: 183–300. water, olivaceous-brown in KOH, 5–15 µm Hawksworth, D. F. & M. S. Cole, 2002. Intrali­ wide, wall up to 2 μm thick; saprobic on bark chen, a new genus for lichenicolous ‘Bispora’ of Quercus, facultatively lichenicolous over and ‘Trimmatostroma’ species. Fungal Diver­ degenerated lichen thalli ...... T. quercicola sity 11: 87–97. 2. Conidial chains indistinct macroscopi- Migula, W., 1921. Fungi imperfecti: Sphaerop- cally, almost entirely blackening large parts sidales, Melanconiales. Kryptogamen-Flora of the host thallus; conidia olivaceous- von Deutschland, Deutsch-Österreich und brown in water, more intensively olivaceous der Schweiz, Band III, 4. Teil, 1. Abteilung. in KOH, 5.5–8.5 µm wide, wall up to 1.2 μm Berlin, Hugo Bermühler Verlag. thick; obligately lichenicolous on lichens Sérusiaux, E., P. Diederich, M. Brand & P. van den with Trentepohlia ...... 3 Boom, 1999. New or interesting lichens and 3. Conidia frequently with a dark outer lichenicolous fungi from Belgium and Lux- wall limited to one side of the conidium embourg. VIII. Lejeunia n. S. 162: 1–95. (resembling conidia of Milospium), wall Sutton, B.C. & K. A. Pirozynski, 1965. Notes on smooth; lichenicolous on cf. Arthonia end­ microfungi. II. Transactions of the British licheri ...... T. hierrense Mycological Society 48: 349–366.

Bull. Soc. Nat. luxemb. 111 (2010) 55