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Notes on three fungicolous fungi: Anastomyces microsporus gen. et sp. nov., Idriella rhododendri sp. nov. and Infundibura adhaerens
Mycol. Res. 101 (11), 1318–1322 (1997)
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Printed in the United Kingdom
Notes on three fungicolous fungi : Anastomyces microsporus gen. et sp. nov., Idriella rhododendri sp. nov. and Infundibura adhaerens
W E N P I N G W U1, 3, B R I AN C. S U T T O N1, 2 A N D A L A N C. G A N GE3 " International Mycological Institute, Bakeham Lane, Egham, Surrey TW20 9TY, U.K. # Old School House, Chapel Street, Bildeston, Suffolk IP7 7EP, U.K. $ School of Biological Sciences, Royal Holloway, University of London, Egham Hill, Egham, Surrey TW20 0EX, U.K.
Three fungicolous hyphomycetes are described and illustrated. Anastomyces gen. nov., with the type species A. microsporus sp. nov., is described on ascomata of Apiospora sp. on dead culms of a grass from China. It differs from comparable genera by its fungicolous habitat, sporodochial conidiomata and polylocal conidiogenous cells mostly with 2 loci. Each locus produces one conidium but two conidia formed from the same conidiogenous cells join together to form H-shaped conidia in the earliest stages of their development. Idriella rhododendri sp. nov. grows over other microfungi and is described from leaf litter of Rhododendron ponticum from England. Infundibura adhaerens was found to be commonly associated with ascomata and conidiomata on dead leaves of R. ponticum in England.
During a general survey of endophytic and epiphytic microfungi from leaves of Rhododendron ponticum L. in southern England, two hyphomycetes growing on fruit bodies of other fungi were collected. These are similar in conidiomatal morphology in that they are sporodochial and produce wet spore masses. One was identified as a new species, Idriella rhododendri, and the other, more frequently collected, was identified as Infundibura adhaerens Nag Raj & W. B. Kendr. (1981) which was described originally from leaf litter. Similarities in fungicolous habit, sporodochial conidiomata with wet spore masses, conidiogenous cells and hyaline aseptate conidia invited comparison between the Idriella species and another fungicolous fungus collected from China by Wenping Wu. However, the unique characters of the latter fungus warrant its description as a new genus and species. Anastomyces microsporus gen. et sp. nov. is described based on conidia which are formed simultaneously in pairs from loci on two conidiogenous cells during which they anastomose forming H-shaped conidia before secession. No descriptions of similar genera were found in a survey of the literature. Descriptive terminology of conidiogenous structures is as recommended by Hennebert & Sutton (1994), whose descriptions of wall relationships and characters of conidiogenous loci are defined and illustrated.
dricae, hyalinae, tenuitunicatae, laeves, multilocales locis duobus in unaquaque cellula conidiogena ; loci apicales, persistentes, cicatricibus clausis et sine fimbriis, protuberantibus, non incrassatis. Conidiogenesis : conidia holoblastica, pariete hologeno, septo uno delimitata, secessione schizolytica. Conidia solitaria, hyalina, aseptata, laevia, tenuitunicata, guttulata, ellipsoidea vel subcylindrica, anastomosantia et mox H-formata duobus conidiis ex isdem cellulis conidiogenis formata. Species typica : Anastomyces microsporus sp. nov.
Conidiomata fungicolous, sporodochial, hyaline to yellow brown, conical, producing wet spore masses ; basal stroma of textura intricata to epidermoidea, composed of hyaline to very pale brown, septate, thin-walled, smooth, branched hyphae. Conidiophores hyaline, septate, branched irregularly, thin-walled, smooth, formed from the stromatic hyphae. Conidiogenous cells integrated, determinate, cylindrical to subcylindrical, hyaline, thin-walled, smooth, multilocal with two loci on each conidiogenous cell ; loci apical, persistent, with the scars closed and without frills, protuberant, unthickened. Conidial ontogeny holoblastic, wall hologenous, delimited by one septum from conidiogenous cells, secession schizolytic. Conidia solitary, hyaline, aseptate, smooth, thin-walled, guttulate, ellipsoid to subcylindrical but soon becoming Hshaped due to anastomosis of two young conidia from the same conidiogenous cell.
Anastomyces W. Wu, B. Sutton & Gange, gen. nov. Conidiomata fungicola, sporodochialia, hyalina vel luteo brunnea, conica, massas conidicas madidas producentia ; stroma basale ex textura intricata vel epidermoidea, ex hyphis hyalinis vel pallidissime brunneis, septatis, tenuitunicatis, laevibus, ramosis compositum. Conidiophora hyalina, septata, irregulariter ramosa, tenuitunicata, laevia, ex hyphis stromatum formata. Cellulae conidiogenae in conidiophoris incorporatae, determinatae, cylindricae vel subcylin-
Anastomyces microsporus W. Wu, B. Sutton & Gange sp. nov. (Figs 1–6) Conidiomata fungicola, sporodochialia, hyalina vel luteo brunnea, conica, 250–350 µm, massas conidicas madidas producentia ; stroma basale ex textura intricata vel epidermoidea, ex hyphis hyalinis vel pallidissime brunneis, septatis, tenuitunicatis, laevibus, ramosis, 2–5 µm crassis compositum. Conidiophora hyalina, septata, irregu-
W. Wu, B. C. Sutton and A. C. Gange
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Figs 1–4. Anastomyces microsporus gen. et sp. nov. Fig. 1. Sporodochia on ascomata of Apiospora sp. (Bar, 2 cm.) Fig. 2. Vertical section of a conidioma on an ascoma. (Bar, 100 µm.) Fig. 3. Textural structure of basal stroma. (Bar, 40 µm.) Fig. 4. Textural structure and conidiophores. lariter ramosa, tenuitunicata, laevia, usque ad 60 µm longa et 2±5–3±5 µm crassa, ex hyphis stromatum formata. Cellulae conidiogenae in conidiophoris incorporatae, determinatae, cylindricae vel subcylindricae, hyalinae, tenuitunicatae, laeves, 6–15¬2±5–4±5 µ, multilocales locis duobus in unaquaque cellula conidiogena ; loci apicales, persistentes, cicatricibus clausis et sine fimbriis, protuberantibus, non incrassatis, usque ad 1 µm lati. Conidia solitaria, hyalina, aseptata, laevia, tenuitunicata, guttulata, ellipsoidea vel subcylindrica, 4–5¬ 2–3 µm, anastomosantia et mox H-formata duobus conidiis ex isdem cellulis conidiogenis formata. Holotypus : In ascomata Apiospora sp. in Gramineae indet., China : Hebei Prov., Chengde, Bishushanzhuang, 18 Nov. 1992, Wenping Wu (W0620), IMI 372457.
Conidiomata fungicolous, sporodochial, hyaline to yellow brown, conical, 250–350 µm diam., producing wet spore masses ; basal stroma of textura intricata to epidermoidea, composed of hyaline to very pale brown, septate, thin-walled, smooth, branched hyphae 2–5 µm wide, basal parts composed of elongated, thick-walled, pale brown cells 5–12 µm wide. Conidiophores hyaline, septate, branched irregularly, thinwalled, smooth, up to 60 µm long, 2±5–3±5 µm wide, formed from stromatic hyphae. Conidiogenous cells integrated, determinate, cylindrical to subcylindrical, hyaline, thin-walled, smooth, 6–15¬2±5–4±5 µm, multilocal and with two loci on each conidiogenous cell, loci apical, persistent, with scars closed and without frills, protuberant, unthickened, up to 1 µm
Figs 5–6. Anastomyces microsporus. Fig. 5. Conidiophores, conidiogenous cells and developing conidia. Note that each locus produces a single conidium simultaneously and anastomosis occurs before secession. Fig. 6. Mature H-shaped conidia. (Bar, 10 µm.)
wide. Conidia solitary, produced in wet spore masses, hyaline, aseptate, smooth, thin-walled, guttulate, ellipsoid to subcylindrical, 4–5¬2–3 µm, with rounded or obtuse ends, but soon becoming H-shaped due to anastomosis of two young conidia from the same conidiogenous cell. Holotype : On ascomata of Apiospora sp. on dead culms of undetermined grass, China : Hebei Province : Chengde, Bishushanzhuang, 18 Nov. 1992, Wenping Wu (W0620), IMI 372457 (holotype). Anastomyces is characterized by its fungicolous habit on ascomata, sporodochial conidiomata with wet spore masses, branched conidiophores and integrated, determinate, multilocal conidiogenous cells, holoblastic, hyaline, aseptate, Hshaped conidia produced by early anastomosis of two young conidia from the same conidiogenous cell. The combination of morphological characters could not be found in any other genus in an extensive survey of the literature. Papilionospora V. Rao & B. Sutton (1975) is the only genus hitherto described with similar anastomosing conidia. Conidia of Papilionospora are also formed simultaneously in pairs and remain joined laterally above the base but they still retain their own walls at the junction area and no anastomosis occurs. Moreover the conidiomata of Papilionospora differ in being synnematous, hyaline, having stipes composed of parallel, unbranched, septate, thick-walled hyphae, and the fungus is not associated with fruiting structures of other fungi. Conidia are dry,
Notes on three fungicolous fungi
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perprolate, smooth, pale brown, thick-walled and eguttulate. In Anastomyces the conidia from each conidiogenous locus anastomose before secession, and the conidial walls at the anastomosis point dissolve resulting in a single propagule. Clinoconidium Pat. (1898) and Botryoconis Syd. & P. Syd. (1906) also produce anastomosing spores but both were incorrectly described as hyphomycetes and actually are basidiomycetes (Malençon, 1953 ; Carmichael et al., 1980). Additionally, the morphology of the spores and fruit bodies produced by these genera differ from those of Anastomyces. Other genera with conidia superficially similar in shape to those of Anastomyces include Araneomyces Ho$ hn., Titaea Sacc., Titaeospora Buba! k, Tricellula Beverw., Triposporina Ho$ hn., Tricladium Ingold, Articulospora Ingold, and Lateriramulosa Matsush. However, in most of these genera, the colonies are dry, the conidiomata are represented by simple conidiophores, and the conidial shape is achieved by branching rather than by two conidia anastomosing (Matsushima, 1975 ; Carmichael et al., 1980 ; Sutton, 1984). Titaea was redescribed and the species revised by Sutton (1984). The four accepted species are fungicolous, especially overgrowing ascomycetes or urediniomycetes. The integrated conidiogenous cells are multilocar often with more than one unthickened denticle, and conidia are holoblastic, hyaline and multicellular with the cells joined by isthmi. Although conidium development and shape appear similar to Anastomyces microsporus each locus of the conidiogenous cell forms a septate conidium which does not anastomose, and only one cell of the multicellular conidium connects to the conidiogenous cell. Other differences include the dry colony and conidiophores which are separate or occasionally loosely aggregated into sporodochial conidiomata. Araneomyces, a monotypic genus with the type species A. acariferus Ho$ hn. described as overgrowing Rosellinia miconiae on leaves of Miconia sp. from the Amazon, was redescribed and illustrated by Sutton (1984). It forms compact, yellowishorange colonies, sporodochial, pulvinate, waxy, yellowishorange-pigmented conidiomata on superficial ascomata of its host. In these characters it is very similar to Anastomyces. However, the integrated conidiogenous cells are monolocar and conidia are holoblastic, hyaline, multicellular, complex, bilaterally symmetrical, consisting of a main axis with two cells each again with two lateral cells growing out from each cell near the septum and orientated in different directions (Sutton, 1984). Anastomosis is a common event in fungi and can occur between different hyphae or between germination tubes produced by different conidia. In the majority of cases in conidia, this occurs normally after secession from conidiogenous cells and during germination. It has been observed frequently in cultures of Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. more than 2 months old. Titaeospora is a coelomycete with conidia often anastomosing in the acervuli by germination tubes produced just above the base of the conidium (Sutton, 1980). However Anastomyces clearly differs
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Figs 7–9. Idriella rhododendri sp. nov. Fig. 7. Sporodochial structure ; note the branched conidiophores and integrated conidiogenous cells. (Bar, 40 µm.) Fig. 8. Conidiophores, conidiogenous cells and developing conidia. (Bar, 20 µm.) Fig. 9. Conidia.
from Titaeospora in conidiomatal structure, conidial morphology and conidiogenesis. Idriella rhododendri W. Wu, B. Sutton & Gange sp. nov. (Figs 7–9) Coloniae luteo aurantiacae, compactae, dispersae. Mycelium partim immersum et partim superficiale, sparsum, ex hyphis hyalinis, septatis, ramosis, laevibus, tenuitunicatis, usque ad 2 µm crassis compositum. Conidiomata sporodochialia, effusa, pulvinata, usque ad 350 µm diam., superficialia in ascomatibus obtecta, dispersa, ceracea, luteo aurantiaca. Conidiophora macronematosa, separata sed in sporodochiis compacta, ad basim et apicem versus irregulariter ramosa, cellula sterili apicali et aliquot cellulis fertilibus intercalaribus, saepe lateraliter et constanter versus praesertim latus orientata, 40–70 µm longa ; pars basalis pallide brunnea, leniter crassitunicata, laevis, 6–12 µm crassa, apicem versus hyalina. Cellulae conidiogenae in conidiophoris incorporatae, intercalares, indeterminatae, multilocales, proliferationibus sympodialibus, lageniformes, 5–8 µm crassae et 6–10 µm longae, regione conidiogena sub septo praecedenti locata ; locis conidiogenis applanatis denticulatis, non-incrassatis et hyalinis. Conidia solitaria, massas madidas producentia, cylindrica vel leniter fusiformia, recta vel leniter curvata, apicem obtusa, basim truncata, tenuitunicata, laevia, guttulata, 12±5–16¬1±5–2 µm. Holotypus : Supra ascomata ignota in foliis emortuis Rhododendri pontici, U.K. : Surrey, Egham, 12 Sep. 1995, Wenping Wu (R214), IMI 372458.
W. Wu, B. C. Sutton and A. C. Gange Colonies yellow-orange, compact, scattered. Mycelium partly immersed and partly superficial, sparse, composed of hyaline, septate, branched, smooth, thin-walled hyphae, up to 2 µm wide. Conidiomata sporodochial, effuse, pulvinate, up to 350 µm diam., overgrowing and superficial on fruit bodies of undetermined ascomycetes, scattered, waxy, yellow-orange in pigmentation. Conidiophores macronematous, separate but also compacted into sporodochia, irregularly branched at the base and above, with a terminal sterile cell and several intercalary fertile cells often orientated laterally and consistently to one side, 40–70 µm long ; basal part pale brown, slightly thickwalled, smooth, 6–12 µm wide, becoming hyaline towards the apex. Conidiogenous cells integrated, intercalary, indeterminate, multilocar, proliferating sympodially, lageniform, 5–8 µm wide, and 6–10 µm long, with the conidiogenous region just below the preceding septum, conidiogenous loci apical, persistent, with scars flattened, denticular, unthickened, unpigmented, closed and without frills. Conidia holoblastic, solitary, wet in mass, cylindrical or slightly fusiform, straight or slightly curved, apex rounded, base truncate, thin-walled, smooth, guttulate, 12±5–16¬1±5–2 µm. Holotype : on ascomata of unknown fungus on dead leaves of Rhododendron ponticum, U.K. : Surrey : Egham, 12 Sep. 1995, Wenping Wu (R214), IMI 372458. Idriella P. E. Nelson & S. Wilh. and Microdochium Syd. are two closely related genera which are distinguished from Dactylaria Sacc. only by conidiogenous cells with nodose apices rather than denticulate apices as seen in Dactylaria (de Hoog, 1985 ; Sutton, Pirozynski & Deighton, 1972 ; von Arx, 1981). The habitat and morphology of species described in Idriella and Microdochium overlap considerably and von Arx (1981) tried to distinguish them based on a series of characters such as habitat, and morphology of chlamydospores, conidiophores, conidiogenous cells and conidia. He suggested Idriella be restricted to saprobic soil-borne or leaf-litter fungi which are dematiaceous and usually with chlamydospores formed in pure culture. The conidiophores of Idriella are either present or absent, conidiogenous cells are either denticulate or apically nodose, and conidia are falcate or lunate. Von Arx (1981) considered species of Microdochium to be parasitic on plants and have white or light colonies in culture ; neither chlamydospores nor erect conidiophores develop. Conidiogenous cells are hyaline with denticles, and the conidia are narrow clavate, obclavate, fusiform or filiform. In these two genera the denticles on conidiogenous cells are not obvious but some species overlap Dactylaria in having slightly denticulate conidiogenous cells. A total of 53 species have been described in Idriella and Microdochium and these differ in morphological characters of conidiophores and conidia (Vittal, 1970 ; Kobayasi, 1971 ; Matsushima, 1971, 1975, 1980, 1985 ; Nicot & Mouchacca, 1971 ; Mouchacca & Samson, 1972 ; Sutton, Pirozynski & Deighton, 1972 ; Sutton & Hodges, 1976 ; de Hoog & Hermanides-Nijhof, 1977 ; Morgan-Jones, 1979 ; von Arx, 1981, 1984 ; Holubova! -Jechova! & Mercado Sierra, 1982 ; Castan4 eda, 1985, 1986 ; Castan4 eda & Arnold, 1985 ; Harris, 1985 ; Galea, Price & Sutton, 1986 ; Samuels, Mu$ ller & Petrini, 1987 ; Subramanian & Bhat, 1987 ; Samuels & Hallett, 1989 ; Castan4 eda & Kendrick, 1991 ; Rodrigues & Samuels, 1992).
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Figs 10–11. Infundibura adhaerens. Fig. 10. Septate sterile hyphae, conidiophores and developing conidia. Fig. 11. Mature conidia with basal and apical appendages. (Bar, 20 µm.)
Species in these three genera are commonly reported as saprobes on leaf litter. Idriella rhododendri grows over other fungi and forms waxy sporodochial conidiomata, and produces irregularly branched conidiophores, multilocar conidiogenous cells and hyaline, aseptate, holoblastic conidia which are very similar to those produced by Anastomyces. However, no anastomoses occur between the conidia either from the same conidiogenous cell or different conidiogenous cells. Though Idriella is not the ideal placement for this fungus, description in Idriella is preferable to the introduction of a new generic name for it. I. rhododendri differs from other Idriella species by the sporodochial conidiomata, wet spore masses, branched conidiophores and conidial morphology. The fungicolous habit and conidial morphology are its main distinguishing characters. Infundibura adhaerens Nag Raj & W. B. Kendr., Can. J. Bot. 59 : 544 (1981). (Figs 10–11) Fungicolous, on fruit bodies of unknown host fungus. Mycelium superficial or partly immersed, pale brown, septate, branched, thin- and smooth-walled, up to 2±5 µm wide. Conidiomata sporodochial, scattered, solitary, superficial, palecoloured, up to 300 µm diam. Setae erect, straight or flexuous, aseptate, cylindrical, pale brown but becoming hyaline towards the apex, base slightly swollen, apex obtuse, wall thin and smooth, arising in same way as conidiogenous cells, up to 100 µm long, 2–2±5 µm wide at the base, 1±5–2 µm at the apex. Conidiophores absent. Conidiogenous cells discrete, de-
Notes on three fungicolous fungi terminate, monolocar, cylindrical, erect, straight or slightly curved at the apex, hyaline, thin-walled and smooth, compactly aggregated, 4–6¬2±5–3±5 µm ; sporulating loci subapical, persistent, with scars closed and without frills, protuberant, unthickened. Conidia produced in pale yellow brown wet spore masses, holoblastic individually hyaline, aseptate, falcate, straight, 18–35¬3–4 µm, apex rounded or obtuse with a funnel-shaped appendage, base with a foot cell and a cytoplasm-free short appendage which is formed by reduction of cytoplasm in one side of the base when conidia are developing. Illustration : Nag Raj & Kendrick (1981). Host plants and fungi : Agathis australis, Picea sitchensis, Pinus sp., conidiomata or ascomata on Rhododendron ponticum. Known distribution : Austria, Canada, New Zealand, U.K. Specimens examined : On dead leaves of Rhododendron ponticum, U.K. : Surrey : Egham, Huntersdale, 10 July 1995, Wenping Wu (R100), IMI 372459 ; 5 Oct. 1995, Wenping Wu (2007a), IMI 372460 ; and also some other collections from the same host in U.K. (in IMI).
Infundibura was described with a single species I. adhaerens by Nag Raj & Kendrick (1981). It is characterized by superficial, sporodochial conidiomata with sterile hyphae, holoblastic conidiogenous cells, and holoblastic–solitary, naviculate conidia measuring 17–35¬3–3±5 µm, with a broad, obtuse apex and a characteristic asymmetric, tapered, foot-like process at the base arising eccentrically at the subapex of conidiogenous cells, apex with a mucoid and infundibuliform appendage with a funnel and a terete stem, base with mucoid and terete appendage. The morphologies of U.K. collections clearly meet the original description. However, the septate character of sterile hyphae was not described by the original authors. Among the four collections examined by them, only one from New Zealand was said to be connected with fruit bodies of ascomycetes or coelomycetes, whereas the colonies in our collection from leaves of R. ponticum are associated with fruit bodies of other fungi (Coleophoma or ?ascomycete). This is the first report of I. adhaerens from the U.K. and of this fungus from any species of Rhododendron worldwide (Farr, Esteban & Palm, 1996). The financial support to Wenping Wu by Novo Nordisk A}S, Denmark, is gratefully acknowledged. REFERENCES Arx, J. A. von (1981). Notes on Microdochium and Idriella. Sydowia 34, 30–38. Arx, J. A. von (1984). Notes on Monographella and Microdochium. Transactions of the British Mycological Society 82, 373–374. Carmichael, J. W., Kendrick, W. B., Conners, I. L. & Sigler, L. (1980). Genera of Hyphomycetes. University of Alberta Press : Alberta, Canada. Castan4 eda Ruiz, R. F. (1985). Deuteromycotina de Cuba. Hyphomycetes II. Instituto de Investigacones Fundamentales en Agricultura Tropical – ‘ Alejandro de Humboldt ’, A. C. C. Castan4 eda Ruiz, R. F. (1986). Fungi Cubenses. Instituto de Investigacones Fundamentales en Agricultura Tropical – ‘ Alejandro de Humboldt ’, A. C. C. (Accepted 14 February 1997)
1322 Castan4 eda Ruiz, R. F. & Arnold, G. R. W. (1985). Deuteromycotina de Cuba. I. Hyphomycetes. Revista del JardıU n BotaU nico Nacional 6, 50–51. Castan4 eda Ruiz, R. F. & Kendrick, W. B. (1991). Ninety-nine conidial fungi from Cuba and three from Canada. University of Waterloo Biological Series 35, 1–132. Farr, D., Bartolome Esteban, H. & Palm, M. E. (1996). Fungi on Rhododendron : A World Reference. Parkway Publishers, Inc. : Boone, North Carolina. Galea, V. J., Price, T. V. & Sutton, B. C. (1986). Taxonomy and biology of the lettuce anthracnose fungus. Transactions of the British Mycological Society 86, 619–628. Harris, D. C. (1985). Microdochium fusarioides sp. nov. from oospores of Phytophthora syringae. Transactions of the British Mycological Society 84, 358–361. Hennebert, G. L. & Sutton, B. C. (1994). Unitary parameters in conidiogenesis. In Ascomycete Systematics : Problems and Perspectives in the Nineties (ed. D. L. Hawksworth), pp. 65–76. Plenum Press : New York. Holubova! -Jechova! , V. & Mercado Sierra, A. (1982). Some new or interesting microfungi from Cuba. Mycotaxon 14(1), 309–315. Hoog, G. S. de (1985). Taxonomy of the Dactylaria complex. IV-VI. Studies in Mycology 26, 1–124. Hoog, G. S. de & Hermanides-Nijhof, E. J. (1977). Survey of black yeasts and allied fungi. Studies in Mycology 15, 178–222. Kobayasi, Y. (ed.) (1971). Mycological reports from New Guinea and the Solomon Islands (1–11). Bulletin of the National Sciences Museum 14(3), 466–467. Malençon, G. (1953). Le Coniodyctium chevalieri Har. et Pat., sa nature et ses affinite! s. Bulletin trimestriel de la SocieU teU Mycologique de France 69, 77–100. Matsushima, T. (1971). Microfungi from the Solomon Islands and Papua–New Guinea. Kobe, published by the author. Matsushima, T. (1975). Icones microfungorum a Matsushima lectorum. Kobe, published by the author. Matsushima, T. (1980). Saprophytic microfungi from Taiwan. Part 1. Hyphomycetes. Matsushima Mycological Memoirs 1, 1–82. Kobe, published by the author. Matsushima, T. (1985). Matsushima Mycological Memoirs 4, 1–68. Kobe, published by the author. Morgan-Jones, G. (1979). Notes on Hyphomycetes. XXX. On three species of Idriella. Mycotaxon 8, 402–410. Mouchacca, J. & Samson, R. A. (1972). Deux nouvelles espe' ces du genre Microdochium Sydow. Revue de Mycologie 37, 267–275. Nag Raj, T. R. & Kendrick, W. B. (1981). Infundibura, a new hyphomycete with unique appendages. Canadian Journal of Botany 59, 542–546. Nicot, J. & Mouchacca, J. (1971). Une nouvelle espe' ce du genre Idriella. Revue de Mycologie 36, 185–193. Patouillard, N. (1898). Champignons nouveaux ou peu connus. Bulletin trimestrie de la SocieU teU Mycologique de France 14, 149–156. Rao, V. & Sutton, B. C. (1975). Synnematous fungi. I. Kavaka 3, 21–26. Rodrigues, K. F. & Samuels, G. J. (1992). Idriella species endophytic in palms. Mycotaxon 43, 271–276. Samuels, G. J. & Hallett, I. C. (1989). Microdochium stoveri and Monographella stoveri, new combinations for Fusarium stoveri and Micronectriella stoveri. Transactions of the British Mycological Society 81, 473–483. Samuels, G. J., Mu$ ller, E. & Petrini, O. (1987). Studies in the Amphisphaeriaceae (sensu lato). 3. New species of Monographella and Pestalosphaeria, and two new genera. Mycotaxon 38, 473–499. Subramanian, C. V. & Bhat, D. J. (1987). Hyphomycetes from South India. I. Some new taxa. Kavaka 15, 41–74. Sutton, B. C. (1980). The Coelomycetes. CMI : Kew. Sutton, B. C. (1984). Notes on Titaea (Hyphomycetes). Transactions of the British Mycological Society 83, 399–413. Sutton, B. C. & Hodges Jr, C. S. (1976). Eucalyptus microfungi : Microdochium and Phaeoisaria species form Brazil. Nova Hedwigia 27, 215–222. Sutton, B. C., Pirozynski, K. A. & Deighton, F. C. (1972). Microdochium. Canadian Journal of Botany 50, 1899–1907. Sydow, H. & Sydow, P. (1906). Novae Fungorum species. III. Annales Mycologici 4, 343–345. Vittal, B. P. R. (1970). A new species of the genus Idriella. Current Science 39, 519–521.
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Notes on three fungicolous fungi : Anastomyces microsporus gen. et sp. nov., Idriella rhododendri sp. nov. and Infundibura adhaerens
W E N P I N G W U1, 3, B R I AN C. S U T T O N1, 2 A N D A L A N C. G A N GE3 " International Mycological Institute, Bakeham Lane, Egham, Surrey TW20 9TY, U.K. # Old School House, Chapel Street, Bildeston, Suffolk IP7 7EP, U.K. $ School of Biological Sciences, Royal Holloway, University of London, Egham Hill, Egham, Surrey TW20 0EX, U.K.
Three fungicolous hyphomycetes are described and illustrated. Anastomyces gen. nov., with the type species A. microsporus sp. nov., is described on ascomata of Apiospora sp. on dead culms of a grass from China. It differs from comparable genera by its fungicolous habitat, sporodochial conidiomata and polylocal conidiogenous cells mostly with 2 loci. Each locus produces one conidium but two conidia formed from the same conidiogenous cells join together to form H-shaped conidia in the earliest stages of their development. Idriella rhododendri sp. nov. grows over other microfungi and is described from leaf litter of Rhododendron ponticum from England. Infundibura adhaerens was found to be commonly associated with ascomata and conidiomata on dead leaves of R. ponticum in England.
During a general survey of endophytic and epiphytic microfungi from leaves of Rhododendron ponticum L. in southern England, two hyphomycetes growing on fruit bodies of other fungi were collected. These are similar in conidiomatal morphology in that they are sporodochial and produce wet spore masses. One was identified as a new species, Idriella rhododendri, and the other, more frequently collected, was identified as Infundibura adhaerens Nag Raj & W. B. Kendr. (1981) which was described originally from leaf litter. Similarities in fungicolous habit, sporodochial conidiomata with wet spore masses, conidiogenous cells and hyaline aseptate conidia invited comparison between the Idriella species and another fungicolous fungus collected from China by Wenping Wu. However, the unique characters of the latter fungus warrant its description as a new genus and species. Anastomyces microsporus gen. et sp. nov. is described based on conidia which are formed simultaneously in pairs from loci on two conidiogenous cells during which they anastomose forming H-shaped conidia before secession. No descriptions of similar genera were found in a survey of the literature. Descriptive terminology of conidiogenous structures is as recommended by Hennebert & Sutton (1994), whose descriptions of wall relationships and characters of conidiogenous loci are defined and illustrated.
dricae, hyalinae, tenuitunicatae, laeves, multilocales locis duobus in unaquaque cellula conidiogena ; loci apicales, persistentes, cicatricibus clausis et sine fimbriis, protuberantibus, non incrassatis. Conidiogenesis : conidia holoblastica, pariete hologeno, septo uno delimitata, secessione schizolytica. Conidia solitaria, hyalina, aseptata, laevia, tenuitunicata, guttulata, ellipsoidea vel subcylindrica, anastomosantia et mox H-formata duobus conidiis ex isdem cellulis conidiogenis formata. Species typica : Anastomyces microsporus sp. nov.
Conidiomata fungicolous, sporodochial, hyaline to yellow brown, conical, producing wet spore masses ; basal stroma of textura intricata to epidermoidea, composed of hyaline to very pale brown, septate, thin-walled, smooth, branched hyphae. Conidiophores hyaline, septate, branched irregularly, thin-walled, smooth, formed from the stromatic hyphae. Conidiogenous cells integrated, determinate, cylindrical to subcylindrical, hyaline, thin-walled, smooth, multilocal with two loci on each conidiogenous cell ; loci apical, persistent, with the scars closed and without frills, protuberant, unthickened. Conidial ontogeny holoblastic, wall hologenous, delimited by one septum from conidiogenous cells, secession schizolytic. Conidia solitary, hyaline, aseptate, smooth, thin-walled, guttulate, ellipsoid to subcylindrical but soon becoming Hshaped due to anastomosis of two young conidia from the same conidiogenous cell.
Anastomyces W. Wu, B. Sutton & Gange, gen. nov. Conidiomata fungicola, sporodochialia, hyalina vel luteo brunnea, conica, massas conidicas madidas producentia ; stroma basale ex textura intricata vel epidermoidea, ex hyphis hyalinis vel pallidissime brunneis, septatis, tenuitunicatis, laevibus, ramosis compositum. Conidiophora hyalina, septata, irregulariter ramosa, tenuitunicata, laevia, ex hyphis stromatum formata. Cellulae conidiogenae in conidiophoris incorporatae, determinatae, cylindricae vel subcylin-
Anastomyces microsporus W. Wu, B. Sutton & Gange sp. nov. (Figs 1–6) Conidiomata fungicola, sporodochialia, hyalina vel luteo brunnea, conica, 250–350 µm, massas conidicas madidas producentia ; stroma basale ex textura intricata vel epidermoidea, ex hyphis hyalinis vel pallidissime brunneis, septatis, tenuitunicatis, laevibus, ramosis, 2–5 µm crassis compositum. Conidiophora hyalina, septata, irregu-
W. Wu, B. C. Sutton and A. C. Gange
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Figs 1–4. Anastomyces microsporus gen. et sp. nov. Fig. 1. Sporodochia on ascomata of Apiospora sp. (Bar, 2 cm.) Fig. 2. Vertical section of a conidioma on an ascoma. (Bar, 100 µm.) Fig. 3. Textural structure of basal stroma. (Bar, 40 µm.) Fig. 4. Textural structure and conidiophores. lariter ramosa, tenuitunicata, laevia, usque ad 60 µm longa et 2±5–3±5 µm crassa, ex hyphis stromatum formata. Cellulae conidiogenae in conidiophoris incorporatae, determinatae, cylindricae vel subcylindricae, hyalinae, tenuitunicatae, laeves, 6–15¬2±5–4±5 µ, multilocales locis duobus in unaquaque cellula conidiogena ; loci apicales, persistentes, cicatricibus clausis et sine fimbriis, protuberantibus, non incrassatis, usque ad 1 µm lati. Conidia solitaria, hyalina, aseptata, laevia, tenuitunicata, guttulata, ellipsoidea vel subcylindrica, 4–5¬ 2–3 µm, anastomosantia et mox H-formata duobus conidiis ex isdem cellulis conidiogenis formata. Holotypus : In ascomata Apiospora sp. in Gramineae indet., China : Hebei Prov., Chengde, Bishushanzhuang, 18 Nov. 1992, Wenping Wu (W0620), IMI 372457.
Conidiomata fungicolous, sporodochial, hyaline to yellow brown, conical, 250–350 µm diam., producing wet spore masses ; basal stroma of textura intricata to epidermoidea, composed of hyaline to very pale brown, septate, thin-walled, smooth, branched hyphae 2–5 µm wide, basal parts composed of elongated, thick-walled, pale brown cells 5–12 µm wide. Conidiophores hyaline, septate, branched irregularly, thinwalled, smooth, up to 60 µm long, 2±5–3±5 µm wide, formed from stromatic hyphae. Conidiogenous cells integrated, determinate, cylindrical to subcylindrical, hyaline, thin-walled, smooth, 6–15¬2±5–4±5 µm, multilocal and with two loci on each conidiogenous cell, loci apical, persistent, with scars closed and without frills, protuberant, unthickened, up to 1 µm
Figs 5–6. Anastomyces microsporus. Fig. 5. Conidiophores, conidiogenous cells and developing conidia. Note that each locus produces a single conidium simultaneously and anastomosis occurs before secession. Fig. 6. Mature H-shaped conidia. (Bar, 10 µm.)
wide. Conidia solitary, produced in wet spore masses, hyaline, aseptate, smooth, thin-walled, guttulate, ellipsoid to subcylindrical, 4–5¬2–3 µm, with rounded or obtuse ends, but soon becoming H-shaped due to anastomosis of two young conidia from the same conidiogenous cell. Holotype : On ascomata of Apiospora sp. on dead culms of undetermined grass, China : Hebei Province : Chengde, Bishushanzhuang, 18 Nov. 1992, Wenping Wu (W0620), IMI 372457 (holotype). Anastomyces is characterized by its fungicolous habit on ascomata, sporodochial conidiomata with wet spore masses, branched conidiophores and integrated, determinate, multilocal conidiogenous cells, holoblastic, hyaline, aseptate, Hshaped conidia produced by early anastomosis of two young conidia from the same conidiogenous cell. The combination of morphological characters could not be found in any other genus in an extensive survey of the literature. Papilionospora V. Rao & B. Sutton (1975) is the only genus hitherto described with similar anastomosing conidia. Conidia of Papilionospora are also formed simultaneously in pairs and remain joined laterally above the base but they still retain their own walls at the junction area and no anastomosis occurs. Moreover the conidiomata of Papilionospora differ in being synnematous, hyaline, having stipes composed of parallel, unbranched, septate, thick-walled hyphae, and the fungus is not associated with fruiting structures of other fungi. Conidia are dry,
Notes on three fungicolous fungi
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perprolate, smooth, pale brown, thick-walled and eguttulate. In Anastomyces the conidia from each conidiogenous locus anastomose before secession, and the conidial walls at the anastomosis point dissolve resulting in a single propagule. Clinoconidium Pat. (1898) and Botryoconis Syd. & P. Syd. (1906) also produce anastomosing spores but both were incorrectly described as hyphomycetes and actually are basidiomycetes (Malençon, 1953 ; Carmichael et al., 1980). Additionally, the morphology of the spores and fruit bodies produced by these genera differ from those of Anastomyces. Other genera with conidia superficially similar in shape to those of Anastomyces include Araneomyces Ho$ hn., Titaea Sacc., Titaeospora Buba! k, Tricellula Beverw., Triposporina Ho$ hn., Tricladium Ingold, Articulospora Ingold, and Lateriramulosa Matsush. However, in most of these genera, the colonies are dry, the conidiomata are represented by simple conidiophores, and the conidial shape is achieved by branching rather than by two conidia anastomosing (Matsushima, 1975 ; Carmichael et al., 1980 ; Sutton, 1984). Titaea was redescribed and the species revised by Sutton (1984). The four accepted species are fungicolous, especially overgrowing ascomycetes or urediniomycetes. The integrated conidiogenous cells are multilocar often with more than one unthickened denticle, and conidia are holoblastic, hyaline and multicellular with the cells joined by isthmi. Although conidium development and shape appear similar to Anastomyces microsporus each locus of the conidiogenous cell forms a septate conidium which does not anastomose, and only one cell of the multicellular conidium connects to the conidiogenous cell. Other differences include the dry colony and conidiophores which are separate or occasionally loosely aggregated into sporodochial conidiomata. Araneomyces, a monotypic genus with the type species A. acariferus Ho$ hn. described as overgrowing Rosellinia miconiae on leaves of Miconia sp. from the Amazon, was redescribed and illustrated by Sutton (1984). It forms compact, yellowishorange colonies, sporodochial, pulvinate, waxy, yellowishorange-pigmented conidiomata on superficial ascomata of its host. In these characters it is very similar to Anastomyces. However, the integrated conidiogenous cells are monolocar and conidia are holoblastic, hyaline, multicellular, complex, bilaterally symmetrical, consisting of a main axis with two cells each again with two lateral cells growing out from each cell near the septum and orientated in different directions (Sutton, 1984). Anastomosis is a common event in fungi and can occur between different hyphae or between germination tubes produced by different conidia. In the majority of cases in conidia, this occurs normally after secession from conidiogenous cells and during germination. It has been observed frequently in cultures of Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. more than 2 months old. Titaeospora is a coelomycete with conidia often anastomosing in the acervuli by germination tubes produced just above the base of the conidium (Sutton, 1980). However Anastomyces clearly differs
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Figs 7–9. Idriella rhododendri sp. nov. Fig. 7. Sporodochial structure ; note the branched conidiophores and integrated conidiogenous cells. (Bar, 40 µm.) Fig. 8. Conidiophores, conidiogenous cells and developing conidia. (Bar, 20 µm.) Fig. 9. Conidia.
from Titaeospora in conidiomatal structure, conidial morphology and conidiogenesis. Idriella rhododendri W. Wu, B. Sutton & Gange sp. nov. (Figs 7–9) Coloniae luteo aurantiacae, compactae, dispersae. Mycelium partim immersum et partim superficiale, sparsum, ex hyphis hyalinis, septatis, ramosis, laevibus, tenuitunicatis, usque ad 2 µm crassis compositum. Conidiomata sporodochialia, effusa, pulvinata, usque ad 350 µm diam., superficialia in ascomatibus obtecta, dispersa, ceracea, luteo aurantiaca. Conidiophora macronematosa, separata sed in sporodochiis compacta, ad basim et apicem versus irregulariter ramosa, cellula sterili apicali et aliquot cellulis fertilibus intercalaribus, saepe lateraliter et constanter versus praesertim latus orientata, 40–70 µm longa ; pars basalis pallide brunnea, leniter crassitunicata, laevis, 6–12 µm crassa, apicem versus hyalina. Cellulae conidiogenae in conidiophoris incorporatae, intercalares, indeterminatae, multilocales, proliferationibus sympodialibus, lageniformes, 5–8 µm crassae et 6–10 µm longae, regione conidiogena sub septo praecedenti locata ; locis conidiogenis applanatis denticulatis, non-incrassatis et hyalinis. Conidia solitaria, massas madidas producentia, cylindrica vel leniter fusiformia, recta vel leniter curvata, apicem obtusa, basim truncata, tenuitunicata, laevia, guttulata, 12±5–16¬1±5–2 µm. Holotypus : Supra ascomata ignota in foliis emortuis Rhododendri pontici, U.K. : Surrey, Egham, 12 Sep. 1995, Wenping Wu (R214), IMI 372458.
W. Wu, B. C. Sutton and A. C. Gange Colonies yellow-orange, compact, scattered. Mycelium partly immersed and partly superficial, sparse, composed of hyaline, septate, branched, smooth, thin-walled hyphae, up to 2 µm wide. Conidiomata sporodochial, effuse, pulvinate, up to 350 µm diam., overgrowing and superficial on fruit bodies of undetermined ascomycetes, scattered, waxy, yellow-orange in pigmentation. Conidiophores macronematous, separate but also compacted into sporodochia, irregularly branched at the base and above, with a terminal sterile cell and several intercalary fertile cells often orientated laterally and consistently to one side, 40–70 µm long ; basal part pale brown, slightly thickwalled, smooth, 6–12 µm wide, becoming hyaline towards the apex. Conidiogenous cells integrated, intercalary, indeterminate, multilocar, proliferating sympodially, lageniform, 5–8 µm wide, and 6–10 µm long, with the conidiogenous region just below the preceding septum, conidiogenous loci apical, persistent, with scars flattened, denticular, unthickened, unpigmented, closed and without frills. Conidia holoblastic, solitary, wet in mass, cylindrical or slightly fusiform, straight or slightly curved, apex rounded, base truncate, thin-walled, smooth, guttulate, 12±5–16¬1±5–2 µm. Holotype : on ascomata of unknown fungus on dead leaves of Rhododendron ponticum, U.K. : Surrey : Egham, 12 Sep. 1995, Wenping Wu (R214), IMI 372458. Idriella P. E. Nelson & S. Wilh. and Microdochium Syd. are two closely related genera which are distinguished from Dactylaria Sacc. only by conidiogenous cells with nodose apices rather than denticulate apices as seen in Dactylaria (de Hoog, 1985 ; Sutton, Pirozynski & Deighton, 1972 ; von Arx, 1981). The habitat and morphology of species described in Idriella and Microdochium overlap considerably and von Arx (1981) tried to distinguish them based on a series of characters such as habitat, and morphology of chlamydospores, conidiophores, conidiogenous cells and conidia. He suggested Idriella be restricted to saprobic soil-borne or leaf-litter fungi which are dematiaceous and usually with chlamydospores formed in pure culture. The conidiophores of Idriella are either present or absent, conidiogenous cells are either denticulate or apically nodose, and conidia are falcate or lunate. Von Arx (1981) considered species of Microdochium to be parasitic on plants and have white or light colonies in culture ; neither chlamydospores nor erect conidiophores develop. Conidiogenous cells are hyaline with denticles, and the conidia are narrow clavate, obclavate, fusiform or filiform. In these two genera the denticles on conidiogenous cells are not obvious but some species overlap Dactylaria in having slightly denticulate conidiogenous cells. A total of 53 species have been described in Idriella and Microdochium and these differ in morphological characters of conidiophores and conidia (Vittal, 1970 ; Kobayasi, 1971 ; Matsushima, 1971, 1975, 1980, 1985 ; Nicot & Mouchacca, 1971 ; Mouchacca & Samson, 1972 ; Sutton, Pirozynski & Deighton, 1972 ; Sutton & Hodges, 1976 ; de Hoog & Hermanides-Nijhof, 1977 ; Morgan-Jones, 1979 ; von Arx, 1981, 1984 ; Holubova! -Jechova! & Mercado Sierra, 1982 ; Castan4 eda, 1985, 1986 ; Castan4 eda & Arnold, 1985 ; Harris, 1985 ; Galea, Price & Sutton, 1986 ; Samuels, Mu$ ller & Petrini, 1987 ; Subramanian & Bhat, 1987 ; Samuels & Hallett, 1989 ; Castan4 eda & Kendrick, 1991 ; Rodrigues & Samuels, 1992).
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Figs 10–11. Infundibura adhaerens. Fig. 10. Septate sterile hyphae, conidiophores and developing conidia. Fig. 11. Mature conidia with basal and apical appendages. (Bar, 20 µm.)
Species in these three genera are commonly reported as saprobes on leaf litter. Idriella rhododendri grows over other fungi and forms waxy sporodochial conidiomata, and produces irregularly branched conidiophores, multilocar conidiogenous cells and hyaline, aseptate, holoblastic conidia which are very similar to those produced by Anastomyces. However, no anastomoses occur between the conidia either from the same conidiogenous cell or different conidiogenous cells. Though Idriella is not the ideal placement for this fungus, description in Idriella is preferable to the introduction of a new generic name for it. I. rhododendri differs from other Idriella species by the sporodochial conidiomata, wet spore masses, branched conidiophores and conidial morphology. The fungicolous habit and conidial morphology are its main distinguishing characters. Infundibura adhaerens Nag Raj & W. B. Kendr., Can. J. Bot. 59 : 544 (1981). (Figs 10–11) Fungicolous, on fruit bodies of unknown host fungus. Mycelium superficial or partly immersed, pale brown, septate, branched, thin- and smooth-walled, up to 2±5 µm wide. Conidiomata sporodochial, scattered, solitary, superficial, palecoloured, up to 300 µm diam. Setae erect, straight or flexuous, aseptate, cylindrical, pale brown but becoming hyaline towards the apex, base slightly swollen, apex obtuse, wall thin and smooth, arising in same way as conidiogenous cells, up to 100 µm long, 2–2±5 µm wide at the base, 1±5–2 µm at the apex. Conidiophores absent. Conidiogenous cells discrete, de-
Notes on three fungicolous fungi terminate, monolocar, cylindrical, erect, straight or slightly curved at the apex, hyaline, thin-walled and smooth, compactly aggregated, 4–6¬2±5–3±5 µm ; sporulating loci subapical, persistent, with scars closed and without frills, protuberant, unthickened. Conidia produced in pale yellow brown wet spore masses, holoblastic individually hyaline, aseptate, falcate, straight, 18–35¬3–4 µm, apex rounded or obtuse with a funnel-shaped appendage, base with a foot cell and a cytoplasm-free short appendage which is formed by reduction of cytoplasm in one side of the base when conidia are developing. Illustration : Nag Raj & Kendrick (1981). Host plants and fungi : Agathis australis, Picea sitchensis, Pinus sp., conidiomata or ascomata on Rhododendron ponticum. Known distribution : Austria, Canada, New Zealand, U.K. Specimens examined : On dead leaves of Rhododendron ponticum, U.K. : Surrey : Egham, Huntersdale, 10 July 1995, Wenping Wu (R100), IMI 372459 ; 5 Oct. 1995, Wenping Wu (2007a), IMI 372460 ; and also some other collections from the same host in U.K. (in IMI).
Infundibura was described with a single species I. adhaerens by Nag Raj & Kendrick (1981). It is characterized by superficial, sporodochial conidiomata with sterile hyphae, holoblastic conidiogenous cells, and holoblastic–solitary, naviculate conidia measuring 17–35¬3–3±5 µm, with a broad, obtuse apex and a characteristic asymmetric, tapered, foot-like process at the base arising eccentrically at the subapex of conidiogenous cells, apex with a mucoid and infundibuliform appendage with a funnel and a terete stem, base with mucoid and terete appendage. The morphologies of U.K. collections clearly meet the original description. However, the septate character of sterile hyphae was not described by the original authors. Among the four collections examined by them, only one from New Zealand was said to be connected with fruit bodies of ascomycetes or coelomycetes, whereas the colonies in our collection from leaves of R. ponticum are associated with fruit bodies of other fungi (Coleophoma or ?ascomycete). This is the first report of I. adhaerens from the U.K. and of this fungus from any species of Rhododendron worldwide (Farr, Esteban & Palm, 1996). The financial support to Wenping Wu by Novo Nordisk A}S, Denmark, is gratefully acknowledged. REFERENCES Arx, J. A. von (1981). Notes on Microdochium and Idriella. Sydowia 34, 30–38. Arx, J. A. von (1984). Notes on Monographella and Microdochium. Transactions of the British Mycological Society 82, 373–374. Carmichael, J. W., Kendrick, W. B., Conners, I. L. & Sigler, L. (1980). Genera of Hyphomycetes. University of Alberta Press : Alberta, Canada. Castan4 eda Ruiz, R. F. (1985). Deuteromycotina de Cuba. Hyphomycetes II. Instituto de Investigacones Fundamentales en Agricultura Tropical – ‘ Alejandro de Humboldt ’, A. C. C. Castan4 eda Ruiz, R. F. (1986). Fungi Cubenses. Instituto de Investigacones Fundamentales en Agricultura Tropical – ‘ Alejandro de Humboldt ’, A. C. C. (Accepted 14 February 1997)
1322 Castan4 eda Ruiz, R. F. & Arnold, G. R. W. (1985). Deuteromycotina de Cuba. I. Hyphomycetes. Revista del JardıU n BotaU nico Nacional 6, 50–51. Castan4 eda Ruiz, R. F. & Kendrick, W. B. (1991). Ninety-nine conidial fungi from Cuba and three from Canada. University of Waterloo Biological Series 35, 1–132. Farr, D., Bartolome Esteban, H. & Palm, M. E. (1996). Fungi on Rhododendron : A World Reference. Parkway Publishers, Inc. : Boone, North Carolina. Galea, V. J., Price, T. V. & Sutton, B. C. (1986). Taxonomy and biology of the lettuce anthracnose fungus. Transactions of the British Mycological Society 86, 619–628. Harris, D. C. (1985). Microdochium fusarioides sp. nov. from oospores of Phytophthora syringae. Transactions of the British Mycological Society 84, 358–361. Hennebert, G. L. & Sutton, B. C. (1994). Unitary parameters in conidiogenesis. In Ascomycete Systematics : Problems and Perspectives in the Nineties (ed. D. L. Hawksworth), pp. 65–76. Plenum Press : New York. Holubova! -Jechova! , V. & Mercado Sierra, A. (1982). Some new or interesting microfungi from Cuba. Mycotaxon 14(1), 309–315. Hoog, G. S. de (1985). Taxonomy of the Dactylaria complex. IV-VI. Studies in Mycology 26, 1–124. Hoog, G. S. de & Hermanides-Nijhof, E. J. (1977). Survey of black yeasts and allied fungi. Studies in Mycology 15, 178–222. Kobayasi, Y. (ed.) (1971). Mycological reports from New Guinea and the Solomon Islands (1–11). Bulletin of the National Sciences Museum 14(3), 466–467. Malençon, G. (1953). Le Coniodyctium chevalieri Har. et Pat., sa nature et ses affinite! s. Bulletin trimestriel de la SocieU teU Mycologique de France 69, 77–100. Matsushima, T. (1971). Microfungi from the Solomon Islands and Papua–New Guinea. Kobe, published by the author. Matsushima, T. (1975). Icones microfungorum a Matsushima lectorum. Kobe, published by the author. Matsushima, T. (1980). Saprophytic microfungi from Taiwan. Part 1. Hyphomycetes. Matsushima Mycological Memoirs 1, 1–82. Kobe, published by the author. Matsushima, T. (1985). Matsushima Mycological Memoirs 4, 1–68. Kobe, published by the author. Morgan-Jones, G. (1979). Notes on Hyphomycetes. XXX. On three species of Idriella. Mycotaxon 8, 402–410. Mouchacca, J. & Samson, R. A. (1972). Deux nouvelles espe' ces du genre Microdochium Sydow. Revue de Mycologie 37, 267–275. Nag Raj, T. R. & Kendrick, W. B. (1981). Infundibura, a new hyphomycete with unique appendages. Canadian Journal of Botany 59, 542–546. Nicot, J. & Mouchacca, J. (1971). Une nouvelle espe' ce du genre Idriella. Revue de Mycologie 36, 185–193. Patouillard, N. (1898). Champignons nouveaux ou peu connus. Bulletin trimestrie de la SocieU teU Mycologique de France 14, 149–156. Rao, V. & Sutton, B. C. (1975). Synnematous fungi. I. Kavaka 3, 21–26. Rodrigues, K. F. & Samuels, G. J. (1992). Idriella species endophytic in palms. Mycotaxon 43, 271–276. Samuels, G. J. & Hallett, I. C. (1989). Microdochium stoveri and Monographella stoveri, new combinations for Fusarium stoveri and Micronectriella stoveri. Transactions of the British Mycological Society 81, 473–483. Samuels, G. J., Mu$ ller, E. & Petrini, O. (1987). Studies in the Amphisphaeriaceae (sensu lato). 3. New species of Monographella and Pestalosphaeria, and two new genera. Mycotaxon 38, 473–499. Subramanian, C. V. & Bhat, D. J. (1987). Hyphomycetes from South India. I. Some new taxa. Kavaka 15, 41–74. Sutton, B. C. (1980). The Coelomycetes. CMI : Kew. Sutton, B. C. (1984). Notes on Titaea (Hyphomycetes). Transactions of the British Mycological Society 83, 399–413. Sutton, B. C. & Hodges Jr, C. S. (1976). Eucalyptus microfungi : Microdochium and Phaeoisaria species form Brazil. Nova Hedwigia 27, 215–222. Sutton, B. C., Pirozynski, K. A. & Deighton, F. C. (1972). Microdochium. Canadian Journal of Botany 50, 1899–1907. Sydow, H. & Sydow, P. (1906). Novae Fungorum species. III. Annales Mycologici 4, 343–345. Vittal, B. P. R. (1970). A new species of the genus Idriella. Current Science 39, 519–521.