Lignicolous freshwater Ascomycota from Thailand: Melanochaeta and Sporoschisma anamorphs

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1 Mycol. Res. 104 (4): (April 2000). Printed in the United Kingdom. 478 Lignicolous freshwater Ascomycota from Thailand: Melanochaeta and Sporoschisma anamorphs Somsak SIVICHAI, Nigel L. HYWEL-JONES and Sayanh SOMRITHIPOL National Center for Genetic Engineering and Biotechnology, National Science and Technology Development Agency, 73 1 Rama VI Road, Rajdhevee, Bangkok 10400, Thailand. Accepted 26 June Two species of Melanochaeta and three species of Sporoschisma are reported from Thailand. Two connections of Sporoschisma and Melanochaeta are confirmed. Melanochaeta hemipsila has Sporoschisma saccardoi as its anamorph with a Chalara synanamorph in culture. Melanochaeta garethjonesii is described as a new species and linked with S. uniseptatum. S. nigroseptatum was recorded but no evidence of a Melanochaeta teleomorph state could be found. INTRODUCTION The poorly known genus Melanochaeta was described from rotten wood in a variety of habitats and currently includes two species. Barr (1990) considered it to be of southern hemispheric distribution based on records at that time and with ascospores having brown mid cells and hyaline end cells much as in Litschaueria. Mu ller et al. (1969) connected M. hemipsila with the dematiaceous Sporoschisma describing S. saccardoi from cultural work. Mu ller & Samuels (1982) demonstrated the relationship between M. aotearoae and S. mirabile, and noted that although the connections between these ascomycetes and their anamorphs has been demonstrated in pure culture, the morphological similarities between the sexual and asexual stages are so great, however, as to render cultural proof of the connection almost superfluous. Significantly, Mu ller & Samuels (1982) were the first to report the production in culture of a Chalara synanamorph. Most recently, Goh et al. (1997) gave a synopsis of Sporoschisma from Hong Kong and noted that although there have been more than thirteen species named there are now only seven accepted species. We are currently studying the colonization of wood in rivers and seeds in the leaf litter of natural forests in Thailand. In the course of this work, three species of Sporoschisma and two species of Melanochaeta were found. These were isolated and connections between Melanochaeta hemipsila, S. saccardoi and the Chalara synanamorph are reported. Melanochaeta garethjonesii is described as a new ascomycete, and is linked with the anamorph S. uniseptatum. In addition, records are provided on the development of S. nigroseptatum which has no known teleomorph. MATERIALS AND METHODS For the study on colonization of woods in rivers the methods used have been described previously (Sivichai et al., 1998a; Sivichai, Hywel-Jones & Jones, 1998 b). For work on seed fungi we collected a variety of seeds in various states of decay from the forest floor. They were cleaned of soil and associated debris and incubated in plastic pots in a humid chamber. Seeds were examined regularly for the presence of fungi, which were then isolated using the same general methods as described by Sivichai et al. (1998a, b). Unless mentioned otherwise, collections were from the Khao Yai National Park. TAXONOMY Melanochaeta hemipsila (Berk. & Broome) E. Mu ll, Harr & Sulmont, Revue de Mycologie 33: 377 (1969). Figs 1 17 Anamorph: Sporoschisma saccardoi E. W. Mason & S. Hughes, Mycological Papers 31: 20 (1949). Colonies effuse, black, hairy, composed of ascomata with mixed tufts of conidiophores (Fig. 1). Ascomata solitary, scattered, superficial, anchored to substrate by an indistinct basal stroma, broadly pyriform, µm high and µm wide, not collapsing when dry; wall smooth and shining, covered with stiff, erect, dark brown to black setae each with a slightly swollen, light brown to colourless cap; up to 300 µm long µm wide. Paraphyses filiform, septate, longer than asci, hyaline. Asci unitunicate, cylindrical to narrow clavate, µm, apex with a refractive, non-amyloid ring, eight-spored (Figs 2 3). Ascospores partially biseriate, fusiform µm with five transverse

2 S. Sivichai, N. L. Hywel-Jones and S. Somrithipol Figs 1 8. Melanochaeta hemipsila and its anamorph Sporoschisma saccardoi. Fig. 1. Ascomata (arrowed) and conidiophores on test block; Fig. 2. Apical apparatus of ascus; Fig. 3. Immature asci; Fig. 4. Ascospores; Figs 5 6. Conidiophores with capitate setae (arrowed); Fig. 7. Chain of conidia; Fig. 8. Conidia. Scale bars: Fig µm; Figs 2 4, 8 10 µm; Figs 5, 6 50 µm. septa (rarely four or six septa), curved, central four cells darker brown and larger than the lighter brown end cells (Fig. 4). Conidiophores with sterile capitate setae arising from a bulbous stroma up to 75 µm diam (Figs 5 6); setae pale brown, paler toward apex, straight or slightly flexuous, one to four septate, µm long and 5 6 µm wide below the terminal swelling which is µm wide and sub-hyaline; conidiophores solitary or in groups of two or three, also arising from stroma, often with one or two, sometimes up to four, capitate setae, µm long, µm at the base, µm wide at the swollen middle part, neck µm wide, dark brown, paler at the torn apex. Conidia formed enteroblastically inside the tubular collarette of the conidiogenous cell, elongate doliiform, µm, fiveseptate, often constricted at the septa the four inner cells dark brown and the two end cells much paler, shorter and somewhat truncate, giving the appearance of constriction between successive conidia of a chain (Fig. 7); the two central cells 7 9 µm long, penultimate cells µm long (Fig. 8). Ascospores and conidia germinating on CMA within 36 h with cool, white fluorescent light at 20, producing a germ tube from either one or both ends or from the middle cell. Colonies on CMA reaching mm diam. in 20 d at 20 in cool, white fluorescent light, diffuse, effuse with hyaline mycelium (Figs 9 11). Mycelium partly immersed, partly superficial, branched, septate, hyphae to 3 µm wide. Sporulation after 20 d profuse throughout the colony, more crowded in the centre. Sporoschisma conidiophores and Chalara state either solitary or arising together with capitate hyphae in scattered fascicles of a few (Figs 12 14). Sporoschisma conidiophores arising with capitate hyphae, macronematous, unbranched, black, smooth, cylindrical, abruptly narrowed at the base µm long, 9 12 µm wide at the base, 9 13 µm wide at the swollen middle part. Conidia catenate, extruded in long chains, µm, not constricted at the septa, five-septate, some three- or four-septate, four inner cells much darker than the two hyaline to pale brown end cells. Chalara conidiophores macronematous, unbranched, pale brown, smooth, tapering gently and uniformly from base to tip, 0 2( 6) septate, 25 35( 150) µm long, µm wide across the venter, narrowing to µm at the collarette (Figs 15 16). Conidia catenate, extruded in long fragmenting chains, µm, cuboid with truncate ends, rarely rectangular, non-septate, sub-hyaline to pale brown (Fig. 17).

3 Melanochaeta and Sporoschisma Figs Sporulation of Melanochaeta hemipsila on CMA. Fig. 9. Colony sporulating after 20 d; Figs Sporoschisma anamorph and Chalara synanamorph; Figs Conidiophores of the Sporoschisma anamorph with capitate setae; Fig. 14. Chalara synanamorph with Sporoschisma anamorph at the base; Figs Chalara synanamorph arising from mycelium; Fig. 17. Chalara conidia. Scale bars: Fig. 9 3 mm; Figs 10, µm; Figs 12, µm; Figs µm. 17 Specimens examined: Teleomorph: Thailand: Nakorn Ratchassima Province, on twig, 25 Sep. 1996, Sivichai (SS109); Prachin Buri Province, stream at km 29 2, on wooden test block (Alstonia scholaris) submerged 4 mo, 8 Jan. 1997, Sivichai (SS244); Prachin Buri Province, stream at km 29 2, on wooden test block (Anisoptera oblonga) submerged 5 mo, 4 Feb. 1997, Sivichai (SS316); Nakorn Ratchassima Province, Mor Singh To Nicola s gibbon research trail, on seed pod of Entada sp. (Leguminosae), 10 Jun. 1998, Somrithipol (SFC274). Anamorph: Thailand: Petchabun Province, Chulaporn Dam, on twig, 1 Oct. 1996, E. B. G. Jones & Sivichai (SS26); Nakorn Ratchassima Province, on twig, 25 Sep. 1996, Sivichai (SS108); Prachin Buri Province, stream at km 29 2, on wooden test block (A. oblonga) submerged for 1 mo, 14 Oct. 1996, Sivichai (SS163); Prachin Buri Province, stream at km 29 2, on wooden test block (A. scholaris) submerged for 2 mo, 14 Nov. 1996, Sivichai (SS166); Prachin Buri Province, stream at km 29 2, on wooden test block (A. scholaris) submerged for 4 mo, 8 Jan. 1997, Sivichai (SS245); Prachin Buri Province, stream at km 29 2, on wooden test block (A. oblonga) submerged for 1 mo, 27 Feb. 1997, Sivichai (SS269); Nakorn Ratchassima Province, on legume pod (Entada sp.), 7 Oct. 1997, Somrithipol (SFC15); Songkhla Province, Prince of Songkhla University, on pod (Hevea brasiliensis), 9 Dec. 1997, Somrithipol (SFC40); Surat Thani Province, Khao Sok National Park, on pod (Hevea brasiliensis), 29 Oct. 1998, Somrithipol (SFC260); Nakorn Ratchassima Province, on legume pod (Entada sp.), 10 Jun. 1998, Somrithipol (SFC274).

4 S. Sivichai, N. L. Hywel-Jones and S. Somrithipol Figs Melanochaeta garethjonesii. Figs Ascomata on test block; Fig. 20. Apical apparatus of ascus; Fig. 21. Squash of asci and paraphyses; Fig. 22. Immature and mature asci; Figs 23. Ascospores. Scale bars: Figs 18, µm; Figs 20, µm; Fig µm; Fig µm. Cultures examined: Isolated from the teleomorph: SS316, SFC274; isolated from the Sporoschisma anamorph: SS26, SS548, SFC16, SFC40 and SFC260. All cultures are stored in the BIOTEC National Culture Collection. Distribution: Teleomorph: Australia (Hyde, pers. comm.), French Guiana, Sri Lanka and Thailand. Anamorph: Australia, Brunei Darussalam, Canada, Ecuador, Hong Kong, Indonesia, Italy, Malaysia, Peru, South Africa, Taiwan and Thailand. Melanochaeta garethjonesii Sivichai & Hywel-Jones, sp. nov. Figs Etym.: In recognition of the support Professor E. B. G. Jones has given to all three authors. Anamorph: Sporoschisma uniseptatum Bhat & W. B. Kendr., Mycotaxon 49: 71 (1993) Ascomata solitaria, dispersa, superficialia, late pyriformia, atrobrunnea, longa, µm crassa statu anamorpho. Peridium nitens, laeve; setis. Setae rigidae, erectae, non-ramosae, atro-brunneae vel atrae, capitatae. Paraphyses filiformes, septatae, hyalinae. Asci cylindrici vel ad anguste clavati, longa, µm crassa, octo-spori, unitunicati. Ascosporae partim biseriatae ad biseriatae, fusiformes, pallidae brunneae, leviter curvae; 4 septa transversa, longa, µm crassa. Coloniae anamorphae effusae, sparsae. Conidiophora mononemata, plerumque 2 5, rigida, erecta vel flexuosa, cylindrica, µm longa µm crassa; inflato medio. Conidia cylindrica, truncata, leviter verruculosa, plerumque 1-septata, raro 2- vel 3-septata, longa, µm crassa, pallentia brunnea. Holotypus: Thailand: Prachin Buri Province, Khao Yai National Park, stream at km 29 2, on wooden test block (A. scholaris) submerged for 11 mo, 12 Aug. 1997, Sivichai SS481 (holotype BIOTEC National Fungus Herbarium). Colonies black, composed of conidiophores and ascomata with capitate setae (Figs 18 19). Ascomata solitary, scattered, superficial, anchored to the substrate by an indistinct basal stroma, broadly pyriform, µm high and µm wide, ascomata not collapsing when dry; wall smooth and shiny, covered with stiff, erect unbranched, dark brown to black setae, to 100 µm long, 5 6 µm wide, capitate, arising from all over surface of ascomatal wall, each with a slightly

5 Melanochaeta and Sporoschisma Figs Sporoschisma uniseptatum. Fig. 24. A representative portion of the colony on test block. Note the long chains of adhering conidia; Fig. 25. Apical portion of phialides; Fig. 26. Conidiophores with basal setae and conidial chain; Fig. 27. Chain of conidia; Fig. 28. Conidia. Scale bars: Fig µm; Figs 25, 27, µm; Fig µm. swollen, light brown to colourless cap. Paraphyses filiform, to 4 µm wide, septate, longer than asci, hyaline (Fig. 21). Asci cylindrical to narrowly clavate µm, unitunicate, apex with a refractive, non-amyloid ring, eightspored (Figs 20 22). Ascospores partially biseriate to biseriate, fusiform µm, predominantly four-septate (rarely three or five), pale brown, slightly curved (Fig. 23). Anamorph colonies effuse, black, sparse (Fig. 24). Conidiophores mononematous, differentiated. Setae present, capitate, erect, straight or flexuous, three to six septate, smooth, pale brown, paler towards the sub-hyaline apex, to 140 µm long, µm wide, with hyaline mucilage around swollen apex; conidiophores mostly in groups of two to five associated with capitate setae, erect, straight or flexuous, cylindrical, µm wide at the base, µm wide at the swollen middle part and µm wide at the venter (Figs 25 26, 32 34). Conidia catenate, cylindrical, truncate at both ends, slightly verruculose, predominantly one-septate, rarely two or three-septate, pale brown, uniform in colour, µm (Figs 27 28, 35). Ascospores germinating on CMA within 36 h with cool, white fluorescent light at 20, producing a germ tube from one or both ends (Fig. 31). Conidia germinating on CMA within 36 h with cool, white fluorescent light at 20, producing a germ tube from one cell or both end cells. Colonies on CMA reaching mm diam in 20 d at 20 in cool, white fluorescent light, diffuse, effuse with hyaline mycelium. Mycelium partly immersed, partly superficial, branched, septate, hyphae to 3 µm wide (Fig. 29). Sporulating after 14 d (Figs 29 30). Specimens examined: Teleomorph: see holotype. No other collection known. Anamorph: Thailand: Prachin Buri Province, stream at km 29 2, on wooden test block (Alstonia scholaris) submerged for 2 mo, 14 Nov. 1996, Sivichai (SS102); Prachin Buri Province, stream at km 29 2, on wooden test block (A. scholaris) submerged for 4 mo, 8 Jan. 1997, Sivichai (SS134); Nakorn Ratchassima Province, stream at Tad Ta Phu, on wooden test block (A. scholaris) submerged for 4 mo, 8 Jan. 1997, Sivichai (SS141); Prachin Buri Province, stream at km 29 2, on wooden test block (A. scholaris) submerged for 5 mo, 4

6 S. Sivichai, N. L. Hywel-Jones and S. Somrithipol Figs Sporulation of Melanochaeta garethjonesii on CMA. Fig. 29. Colony sporulating after 14 d; Fig. 30. Sporulation; Fig. 31. Germination of ascospores on CMA after 36 h; Figs Squash of Sporoschisma conidiophores; Fig. 35. Conidia. Scale bars: Fig mm; Fig mm; Figs µm; Fig µm. Feb. 1997, Sivichai (SS193); Prachin Buri Province, stream at km 29 2, on wooden test block (A. oblonga) submerged for 5 mo, 4 Feb. 1997, Sivichai (SS202); Nakorn Ratchassima Province, stream at Tad Ta Phu, on wooden test block (A. scholaris) submerged for 5 mo, 4 Feb. 1997, Sivichai (SS208); Prachin Buri Province, stream at km 29 2, on wooden test block (A. scholaris) submerged for 6 mo, 27 Feb. 1997, Sivichai (SS263); Prachin Buri Province, stream at km 29 2, on wooden test block (A. oblonga) submerged for 4 mo, 8 Jan. 1997,

7 Melanochaeta and Sporoschisma Figs Sporoschisma nigroseptatum: Fig. 36. A representative portion of the colony on test block; Fig. 37. Apical portion of phialides; Fig. 38. Conidiophores and setae; Fig. 39. Conidia. Scale bars: Fig mm; Figs 37, µm; Fig µm. Sivichai, (SS312); Nakorn Ratchassima Province, stream at Tad Ta Phu, on wooden test block (A. scholaris) submerged for 7 mo, 10 Apr. 1997, Sivichai (SS334); Prachin Buri Province, stream at km 29 2, on wooden test block (A. scholaris) submerged for 11 mo, 12 Aug. 1997, Sivichai (SS482); Nakorn Ratchassima Province, stream at Tad Ta Phu, on wooden test block (A. scholaris) submerged for 11 mo, 12 Aug. 1997, Sivichai (SS483); Satun Province, Thaleban National Park, on legume pod, 11 Dec. 1997, Somrithipol (SFC 127); Songkhla Province, Ton Nga Chang wildlife Sanctuary, on legume pod, 13 Nov. 1997, Somrithipol (SFC 172). Cultures examined: Isolated from the teleomorph: SS481; isolated from the Sporoschisma anamorph: SS102, SS134, SS141, SS481 ( SS482), SS483, SFC127 and SFC172. All cultures are stored in the BIOTEC National Culture Collection. Distribution: Teleomorph: Thailand, type locality. Anamorph: India, Seychelles and Thailand. Sporoschisma nigroseptatum D. Rao & Rao, Mycopathologia et Mycologia Applicata 24: 82 (1964). Figs Teleomorph: unknown. Colonies effuse, black, hairy (Fig. 36). Setae sparse, capitate, erect, straight or flexuous, 0 3 septate, smooth, pale brown, paler towards the sub-hyaline apex, up to 150 µm long, 5 6 µm wide, sometimes proliferating once percurrently; swollen apex 8 10 µm wide, surrounded by mucilage. Conidiophores mainly solitary or in groups of two to three, each with zero to two setae, erect, straight or flexuous, cylindrical, µm long, µm wide at the base, µm wide at the swollen middle part, thick-walled, smooth, dark brown (Figs 37 38). Conidiogenous cells monophialidic, terminal, integrated, lageniform, consisting of a slightly swollen venter µm wide, frayed at the apex. Conidia catenate, elongate doliiform, sub-truncate at both ends, µm, five septate, with four inner cells dark brown and the two end cells hyaline to pale brown, septa thick, the inner three septa µm thick, the two distal septa ca µm thick, the two central cells ( µm) are longer than the penultimate cells (5 6 µm) (Fig. 39). Conidia germinating on CMA within 48 h with cool, white fluorescent light at 20, producing a germ tube from both end cells or from the middle cells. Colonies on CMA reaching mm diam. in 30 d at 20 in cool, white fluorescent light, diffuse, effuse, hyaline mycelium with aerial hyphae. Mycelium partly immersed, partly superficial, branched, septate, hyphae up to 3 µm wide; colony becoming black with a ring of sporulation after 14 d, conidiophores more crowded in the middle. Specimens examined: Thailand: Nakorn Ratchassima Province, stream at Tad Ta Phu, on wooden test block (A. oblonga) submerged for 4 mo, 8 Jan. 1997, Sivichai (SS311); Nakorn Ratchassima Province, stream at Tad Ta Phu, on wooden test block (A. scholaris) submerged for 8 mo, 6 May 1997, Sivichai (SS485). Cultures examined: Isolated from the anamorph: SS311 and SS485, deposited in the BIOTEC National Culture Collection. Distribution: Australia, Hong Kong, India, Japan, New Zealand and Thailand.

8 S. Sivichai, N. L. Hywel-Jones and S. Somrithipol 485 Table 1. Species of Sporoschisma and their known teleomorphs. Teleomorph Sporoschisma anamorph Chalara synanamorph Ascomata (µm) Asci (µm) Ascospores (µm) Ascospore septa M. hemipsila S. saccardoi Present M. aotearoae S. mirabile Present M. garethjonesii S. uniseptatum Absent DISCUSSION Melanochaeta garethjonesii has larger ascomata and bigger asci than M. aotearoae (Table 1). Although ascomata, asci and ascospores have similar sizes to those of M. hemipsila, there are significant differences in the ascospore colour and number of septa (Table 1). Importantly the ascospores of M. garethjonesii are a uniform brown whereas those of M. hemipsila have pale end cells. The most significant difference between the three species, is in the anamorph. Although the Sporoschisma state of M. hemipsila appears to have a pan-global distribution the teleomorph is known only from a few scattered tropical or subtropical records. Furthermore, these records are mostly from the Asia Pacific region, although it has been reported from French Guiana (Courtecuisse et al., 1996). Mu ller & Samuels (1982) note that Sporoschisma is characterised by cylindrical, monophialidic conidiophores; integrated, tubular phialides; pale brown to nearly black, cylindrical, catenate, phragmoconidia and erect setae, each of which has a somewhat swollen tip that is enclosed with mucilaginous material. Mu ller & Samuels (1982) found that Melanochaeta aotearoae is synanamorphic, producing a Sporoschisma with a Chalara state in culture. Nag Raj & Kendrick (1975) discussed the pleomorphic nature of Chalara and related species, but discussed this with respect to the formation of chlamydospores. Mu ller & Samuels (1982) noted that no species of Sporoschisma is known to be polymorphic with the possible exception of S. juvenile. In our study we report the first association of a Chalara anamorph with S. saccardoi. Significantly, isolations made from the Sporoschisma state failed to produce the Chalara in culture while isolations from ascospores of the Melanochaeta produced both anamorph states. Although we could produce S. uniseptatum in culture from M. garethjonesii, this failed to produce a Chalara synanamorph. For now, we have no explanation for these observations. Melanochaeta hemipsila and Sporoschisma saccardoi were both recorded from hard and soft woods. Both states were, however, only recorded from the seasonal stream at km 29 2 and not from Tad Tha Phu (which flows all through the year). M. hemipsila was recorded from test blocks that had been submerged for 4 5 mo, which coincided with the end of the rainy season and the start of the cool dry season when river levels begin to fall. In contrast, S. saccardoi also appeared on wood that had only been submerged for 1 or 2 mo. Melanochaeta garethjonesii was only recorded once from km 29 2 on soft wood that had been submerged for 11 mo and would appear to be rare. In contrast, its anamorph S. uniseptatum was the most commonly encountered Sporoschisma in our study and was recorded within two months of test blocks being submerged. That there are many records of Sporoschisma compared with the few records for the Melanochaeta teleomorphs suggest that conditions necessary for the development of the Melanochaeta are not often met in the field. It is possible the appearance of the teleomorph over time is due to the increased chance of different genetic states colonizing the wood substrate. Without compatible crosses the teleomorph would not be expressed. Time will increase the chances of fertile crosses coming together if these are present in the stream to begin with. We are finding increasing numbers of homothallic ascomycetes in our studies (Hywel-Jones & Sivichai, unpubl. obs.). The circumstantial evidence of teleomorph rarity compared with anamorph prevalence, however, and the late appearance of the teleomorph compared with the anamorph, leads us to conclude that Melanochaeta could be heterothallic. This might also account for the Chalara state only appearing in cultures originating from the ascospores. We did not note germination of the Chalara state and do not know what role this might play in the life-cycle in nature. ACKNOWLEDGEMENTS Professor E. B. Gareth Jones is thanked for the guidance that he has given to this work. Dr Kevin D. Hyde is thanked for his support and for the provision of unpublished data. This work was supported by the TRF BIOTEC Special Programme for Biodiversity Research and Training grant BRT REFERENCES Barr, M. E. (1990). Prodromus to nonlichenized, pyrenomycetous members of Class Hymenoascomycetes. Mycotaxon 39, Courtecuisse, R., Samuels, G. J., Hoff, M., Rossman, A. Y., Cremers, G., Huhndorf, S. M. & Stephenson, S. L. (1996). Check-list of fungi from French Guiana. Mycotoxon 57, Goh, T. K., Ho, W. H., Hyde, K. D. & Umali, T. M. (1997). New records and species of Sporoschisma and Sporoschismopsis from submerged wood in the tropics. Mycological Research 101, Mu ller, E. & Samuels, G. J. (1982). Anamorphs of pyrenomycetous ascomycetes. III. The Sporoschisma and Chalara anamorphs of Melanochaeta aotearoae. Sydowia 35, Mu ller, E., Harr, J. & Sulmont, P. (1969). Deux Ascomycetes dont le stade conidien presente des conidies phaeophragmie es (endoge nes). Revue de Mycologie 33, Nag Raj, T. R. & Kendrick, B. (1975). A Monograph of Chalara and allied Genera. Wilfrid Laurier University Press: Waterloo, Ontario, Canada. Sivichai, S., Goh, T.-K., Hyde, K. D. & Hywel-Jones, N. L. (1998a). The genus Brachydesmiella from submerged wood in the tropics, including a new species and a new combination. Mycoscience 39, Sivichai, S., Hywel-Jones, N. L. & Jones, E. B. G. (1998b). Lignicolous freshwater Ascomycota from Thailand: 1. Ascotaiwania sawada and its anamorph state Monotosporella. Mycoscience 39, Corresponding Editor: D. L. Hawksworth