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A revision of Dactylaria, with description of D. tunicata sp. nov. from submerged wood in Australia
Mycol. Res. 101 (10) : 1265–1272 (1997)
1265
Printed in the United Kingdom
A revision of Dactylaria, with description of D. tunicata sp. nov. from submerged wood in Australia
T. K. G O H A N D K. D. H Y D E Department of Ecology and Biodiversity, The University of Hong Kong, Pokfulam Road, Hong Kong
Dactylaria tunicata sp. nov. from submerged wood in a freshwater stream from Australia is described and illustrated. It differs from all previously described species of Dactylaria in having uniseptate conidia with a hyaline sheath. The 41 species of Dactylaria accepted by de Hoog are listed, together with a further 41 validly published species of Dactylaria post de Hoog. A taxonomic key to 37 of the 41 post de Hoog species is provided, based on the literature, together with a composite illustration of their conidial morphology. The other four species are discussed, but they are not considered to be Dactylaria species (sensu de Hoog).
Since the establishment of Dactylaria Sacc. (1880), based on the lectotype species D. purpurella (Sacc.) Sacc., there have been numerous additions to the genus. Hyphomycetes which produce hyaline, thin-walled, 1–several septate conidia on sympodially proliferating, erect, hyaline or brown conidiophores, with more or less cylindrical denticles, are usually placed in the genus. These include lignicolous saprotroph, foliicolous plant pathogens, animal disease agents, fungicolous hyperparasites and nematophagous species. The generic concept has become confused, taxonomically and ecologically heterogeneous, though species distinction is somewhat easier. The heterogeneity of Dactylaria and similar fungi has provoked some early generic revision (Bhatt & Kendrick, 1968 ; Rifai, 1968 ; de Hoog & von Arx, 1973 ; Schenck, Kendrick & Pramer, 1977). Recent taxonomic revision of the Dactylaria-complex is given by de Hoog & Hennebert (1983), de Hoog, van Oorschot & Hijwegen (1983), de Hoog (1985), van Oorschot (1985), and de Hoog & van Oorschot (1985). In these publications, species disposed in Dactylaria (sensu lato) have been reidentified and reevaluated. The genus was redefined and subdivided into four sections (de Hoog, 1985), i.e. Dactylaria, Diplorhinotrichum (Ho$ hn.) de Hoog, Mirandina (G. Arnaud ex Matsush.) de Hoog, and Pleurophragmium (Costantin) de Hoog, with 37 allied genera, such as Arthrobotrys Corda, Duddingtonia R. C. Cooke, Geniculifera Rifai, Neta Shearer & J. L. Crane, Scolecobasidium E. V. Abbott, and Subulispora Tubaki. Fortyone species were accepted in Dactylaria (sensu de Hoog) and keys were given to groups of species in each section (de Hoog, 1985). Keys and an annotated check-list of published epithets were also provided to species of Neta, Subulispora and other allied genera of the Dactylaria-complex (de Hoog, 1985 ; de Hoog & van Oorschot, 1985). When grown in different culture media, many species of the
Dactylaria-complex exhibit morphological variations. In his review, de Hoog (1985) concluded that the taxonomy of these fungi is still in an exploratory phase and awaits a balanced understanding of their biology and variability. Although four sections were proposed to express the gross morphological differences within Dactylaria, de Hoog regarded the generic delimitation at present as artificial. Despite such generic revision by de Hoog, Matsushima (1987, 1989, 1993, 1995) still maintained Mirandina and Pleurophragmium as genera separate from Dactylaria (sensu de Hoog), describing several new species. De Hoog’s generic revision, with the acceptance of 41 species in Dactylaria (de Hoog, 1985 ; Table 1), represents a major contribution to fungal taxonomy. During the past decade, a further 40 species of Dactylaria (sensu de Hoog) have been validly published (Dixon & Salkin, 1986 ; Arambarri, Cabello & Mengascini, 1987 ; Dixon et al., 1987 ; Matsushima, 1987, 1989, 1993 ; Salkin & Dixon, 1987 ; Castan4 eda Ruiz, 1988 ; Castan4 eda Ruiz & Kendrick, 1990 ; Cazau, Arambarri & Cabello, 1990 ; Castan4 eda Ruiz & Kendrick, 1991 ; Tzean & Chen, 1991 ; Upadhyay & Mankau, 1991 ; Vittal & Dorai, 1991 ; Braun & Crous, 1992 ; Glockling, 1992 ; Jiao, Liu & Wang, 1994 ; Wu, Sutton & Gange, 1996). Most of the species are described from decaying plant material ; however, D. endophytica W. Wu, B. Sutton & Gange (1996) occurs as an endophyte from a living symptomless leaf of Prunus lusitanica. Dactylaria constricta (E. V. Abbott) D. M. Dixon & Salkin var. constricta, and the other variety D. constricta var. gallopava (Ivimey Cook) Salkin & D. M. Dixon (1987) are reported to be pathogens of birds and other animals, including man. Most species of Dactylaria (sensu lato) produce smooth, hyaline conidia which are usually fusiform, naviculate, or cylindrical in shape, but in some species they may be obpyriform, obclavate, fabiform, botuliform, or other shapes.
Revision of Dactylaria and one new species Table 1. Forty-one species of Dactylaria accepted by De Hoog Section Dactylaria Dactylaria acerosa Matsush., Icones Microfungorum a Matsushima Lectorum, p. 48 (1975). Dactylaria clavata Matsush., Icones Microfungorum a Matsushima Lectorum, p. 49 (1975). Dactylaria dimorphospora Veenb.-Rijks, Acta Botanica Neerlandica 22, 641 (1973). Dactylaria humicola G. C. Bhatt & W. B. Kendr. sensu Knauss & Alfieri, Proceedings of Florida States Horticultural Society 83, 441 (1970). 3 Dactylaria humicola G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). ¯ Dactylaria pseudoampulliformis Matsush., Icones Microfungorum a Matsushima Lectorum, p. 52 (1975). Dactylaria lanosa Malla & W. Gams, Persoonia 6, 193 (1971). Dactylaria ponapensis Matsush., Matsushima Mycological Memoirs 4, 7 (1985). Dactylaria purpurella (Sacc.) Sacc., Michelia 2, 20 (1880) ; Sylloge fungorum omnium hucusque cognitorum 4, 195 (1886). 3 Acrothecium purpurellum Sacc., Michelia 1, 75 (1877). Dactylaria scolecospora P. M. Kirk, Transactions of the British Mycological Society 81, 404 (1983). Section Diplorhinotrichum Dactylaria affinis (O. Rostr.) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). 3 Diplorhinotrichum affine O. Rostr., Dansk Botanisk Arkiv 2, 43 (1916). Dactylaria africana (S. Hughes) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). 3 Diplorhinotrichum africanum S. Hughes, Mycological Papers 47, 9 (1951). Dactylaria biseptata Matsush., Icones Microfungorum a Matsushima Lectorum, p. 48 (1975). Dactylaria candibula (Ho$ hn.) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). 3 Diplorhinotrichum candidulum Ho$ hn., Sitzungsberichte Akademie Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse Abteilung 111, 1040 (1902). Dactylaria chrysosperma (Sacc.) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1257 (1968). 3 Rhinotrichum chrysospermum Sacc., Nuovo Giornale Botanico Italiano 8, 196 (1876). 3 Didymotrichum chrysospermum (Sacc.) Ho$ hn., Sitzungsberichte Akademie Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse Abteilung 123, 140 (1914). 3 Diplorhinotrichum chrysospermum (Sacc.) S. Hughes, Canadian Journal of Botany 36, 764 (1958). Dactylaria curviclavata Matsush., Matsushima Mycological Memoirs 3, 7 (1983). Dactylaria fulva R. Y. Roy & Gujarati, Lloydia 28, 53 (1965). ¯ Dactylaria indica Agarwal & Chowdhry, Current Science 46, 352 (1977). Dactylaria fusifera (Berk. & M. A. Curtis) de Hoog, Studies in Mycology 26, 17 (1985). 3 Rhinotrichum fusiferum Berk. & M. A. Curtis, Grevillea 3, 108 (1875). 3 Didymotrichum fusiferum (Berk. & M. A. Curtis) Linder, Lloydia 5, 207 (1942). Dactylaria inaequilatera Matsush., Matsushima Mycological Memoirs 3, 8 (1983). Dactylaria intermedia Matsush., Matsushima Mycological Memoirs 1, 25 (1980). Dactylaria naviculiformis Matsush., Icones Microfungorum a Matsushima Lectorum, p. 51 (1975). Dactylaria subuliphora Matsush., Icones Microfungorum a Matsushima Lectorum, p. 52 (1975). Dactylaria uniseptata Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 19 (1971).
1266 Section Pleurophragmium Dactylaria acerina Matsush., Matsushima Mycological Memoirs 3, 7 (1983). Dactylaria cymbiformis Matsush., Matsushima Mycological Memoirs 1, 25 (1980). Dactylaria dimorpha Matsush., Icones Microfungorum a Matsushima Lectorum, p. 49 (1975). Dactylaria dioscorae M. B. Ellis, More Dematiaceous Hyphomycetes, p. 171 (1976). Dactylaria fusarioidea Matsush., Icones Microfungorum a Matsushima Lectorum, p. 50 (1975). Dactylaria madrasensis Matsush., Matsushima Mycological Memoirs 3, 8 (1983). Dactylaria mitrata Matsush., Matsushima Mycological Memoirs 3, 8 (Sep. 1983). ¯ Hemibeltrania mitrata P. M. Kirk, Mycotaxon 18, 275 (Oct. 1983). Dactylaria mucronulata Ellis & Langl., Journal of Mycology 6, 35 (1890). Dactylaria kumamotoensis Matsush., Matsushima Mycological Memoirs 2, 5 (1981). Dactylaria parvispora (Preuss) de Hoog & Arx, Kavaka 1, 58 (1973). 3 Cordana parvispora Preuss, Linnaea 25, 728 (1852). 3 Pleurophragmium parvisporum (Preuss) Hol.-Jech., CeskaU Mykologie 36, 223 (1972). ¯ Pleurophragmium simplex (Berk. & Broome) S. Hughes, Canadian Journal of Botany 36, 798 (1958). 3 Acrothecium simplex Berk. & Broome, Annals and Magazine of Natural History III 7, 382 (1861). ¯ Acrothecium simplex Berk. & Broome var. elatum Grove, Journal of Botany, London 24, 203 (1886). ¯ Pleurophragmium bicolor Costantin, Les MuceU dineU eU s, p. 100 (1888). Section Mirandina Dactylaria breviphora Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 37 (1971). Dactylaria congregata de Hoog, Studies in Mycology 26, 30 (1985). Dactylaria fragilis de Hoog, Studies in Mycology 26, 30 (1985). Dactylaria fusiformis Shearer & J. L. Crane, Mycologia 63, 243 (March, 1971) ; non Dactylaria fusiformis (Matsush.) de Hoog & Arx, Kavaka 1, 58 (1973). ¯ Mirandina typica Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 38 (Sep. 1971). Dactylaria irregularis de Hoog, Studies in Mycology 26, 32 (1985). Dactylaria lepida Minter, Transactions of the British Mycological Society 74, 570 (1980). Dactylaria obtriangularia Matsush., Icones Microfungorum a Matsushima Lectorum, p. 51 (1975). Doubtful species Mirandina corticola G. Arnaud ex Matsush., Icones Microfungorum a Matsushima Lectorum, p. 96 (1975). 3 Mirandina corticola G. Arnaud, Bulletin de la SocieU teU Mycologique de France 68, 206 (1952) (nomen nudum sine descriptione latina : Article 36, ICBN). ¯ Mirandina yushanensis Matsush., Matsushima Mycological Memoirs 3, 12 (1983). The range of variability of this species is not certain. The shapes and sizes of the conidiophores, denticles and conidia were found to have slight differences based on materials on natural substrate and in culture. Mirandina cylindrospora Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 37 (1971). This species resembles D. biseptata and D. fulva, differing in having conidia with up to 3 septa. Mirandina dactylelloides Matsush., Matsushima Mycological Memoirs 1, 50 (1970). This species is closer to D. dimorphospora, but, it also resembles Trichoconis echinophilum (C. Massal.) de Hoog, Fusarium pallidoroseum (Cooke) Sacc. (Pseudohansfordia venezuelensis (Deighton & Piroz.) de Hoog), and species of Vermispora Deighton & Piroz.
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Figs 1–5. Dactylaria tunicata. Figs 1–2. Squash mounts of conidiophores and conidia in lactophenol. Figs 3–5. Conidia. Note the hyaline gelatinous sheath around the conidia. Scale bars : 1, 50 µm ; 2, 20 µm ; 3–5, 10 µm.
Dactylaria appendiculata Cazau, Arambarri & Cabello (1990) and D. argentina (Aramb. & Mengasc.) Cabello & Cazau (Cazau et al., 1990) produce conidia with an appendiculate apex or filiform extension of the apical cell. Dactylaria isthmospora Cabello, Mengasc. & Aramb. (Arambarri et al., 1987) is somewhat atypical because its conidiophores bear 3–5 terminal and intercalary denticles which are arranged in whorls, and the conidia are somewhat clavate with a globose apex connected by an isthmus. Dactylaria laxa R. F. Castan4 eda (1988) is another atypical species which produces two-celled, fusiform, greyish, verrucose conidia from conidiphores which are pigmented, multiseptate and geniculate, with terminal and intercalary, chiefly monodenticulate conidiogenous cells. It would be more suitable to place this species in Scolecobasidium E. V. Abbott or Ochroconis de Hoog & Arx (1973). During our investigation of freshwater microfungi occurring on submerged woody material from streams in Queensland, Australia, we found a hyphomycete representing a member of the Dactylaria-complex which we consider to be new to science. In this paper, we describe this species as Dactylaria tunicata sp. nov. (Figs 1–7). It is unique amongst Dactylaria species in having navicular, hyaline, 1-septate conidia with a hyaline gelatinous sheath, and belongs to the section Diplorhinotrichum (Ho$ hn.) de Hoog (1985) as it produces pigmented conidiophores with conspicuous denticles at the apical region. With the description of this new hyphomycete in Dactylaria, there are presently 82 accepted species in the genus. A list of all 41 Dactylaria species which have been validly published post de Hoog (1985) is given in Table 2. In this paper, we do not attempt to reclassify each species according to the emended generic concept of Dactylaria (sensu de Hoog), although we realized that the genus is heterogeneous. A taxonomic key to 37 of those species growing on plant material (published post de Hoog, 1985) is given here
Figs 6–7. Dactylaria tunicata, diagramatic representation. Fig. 6. Conidiophores bearing conidia. Fig. 7. Conidia with gelatinous sheath. Scale bar, 10 µm.
based on the literature, and a composite illustration of their conidial morphology is provided (Figs 8–44). The atypical species D. isthmospora, D. laxa, as well as the two varieties of D. constricta (not foliicolous or lignicolous species) are excluded as we do not believe these are true Dactylaria species (sensu de Hoog). Since de Hoog (1985) has already provided
Revision of Dactylaria and one new species Table 2. Species of Dactylaria published post de Hoog, 1985 ; de Hoog & van Oorschot, 1985 Dactylaria appendiculata Cazau, Aramb. & Cabello, Mycotaxon 38, 21 (1990). Dactylaria arecae (Matsush.) R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 26 (1991). 3 Pleurophragmium arecae Matsush., Matsushima Mycological Memoirs 5, 23 (1987). Dactylaria argentina (Aramb. & Mengasc.) Cabello & Cazau, Mycotaxon 38, 22 (1990). 3 Subulispora argentina Aramb. & Mengasc., Mycotaxon 30, 264 (1987). Dactylaria aspirensis R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 26 (1991). Dactylaria attenuata Matsush., Matsushima Mycological Memoirs 5, 10 (1987). Dactylaria botulispora R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 32, 13 (1990). Dactylaria constricta (E. V. Abbott) D. M. Dixon & Salkin var. constricta, Mycotaxon 29, 379 (1987). 3 Scolecobasidium constrictum E. V. Abbott, Mycologia 19, 30 (1927). Dactylaria constricta var. gallopava (Ivimey Cook) Salkin & D. M. Dixon, Mycotaxon 29, 379 (1987). 3 Diplorhinotrichum gallopavum Ivimey Cook, Sabouraudia 3, 241 (1964). Dactylaria cubensis R. F. Castan4 eda, Fungi Cubensis III, 4 (1988). Dactylaria cylindrospora (Matsush.) R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 27 (1991). 3 Pleurophragmium cylindrosporum Matsush., Icones Microfungorum a Matsushima Lectorum, p. 115 (1975). Dactylaria endophytica W. Wu, B. Sutton & Gange, Mycological Research 100, 524 (1996). Dactylaria eucalypti Vittal & Dorai, Mycological Research 95, 501 (1991). Dactylaria fabiformis Matsush., Matsushima Mycological Memoirs 7, 48 (1993). Dactylaria fecundissima R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 27 (1991). Dactylaria havanensis R. F. Castan4 eda, Fungi Cubenses III, 4 (1988). Dactylaria hemibeltranioidea R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 27 (1991). Dactylaria hoogii R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 28 (1991). Dactylaria iriomoteana Matsush., Matsushima Mycological Memoirs 5, 10 (1987). Dactylaria isoscelispora W. B. Kendr. & R. F. Castan4 eda, University of Waterloo Biology Series 32, 16 (1990). Dactylaria isthmospora Cabello, Mengasc. & Aramb., Mycotaxon 30, 263 (1987). Dactylaria laxa R. F. Castan4 eda, Fungi Cubenses III, 5 (1988). Dactylaria leptosphaeriicola U. Braun & Crous, Mycotaxon 45, 101 (1992). Dactylaria lignicola W. B. Kendr. & R. F. Castan4 eda, University of Waterloo Biology Series 35, 29 (1991). Dactylaria longidentata Cazau, Aramb. & Cabello, Mycotaxon 38, 22 (1990). Dactylaria longispora Matsush., Matsushima Mycological Memoirs 5, 10 (1987). Dactylaria lunata Tzean & J. L. Chen, Mycological Research 95, 1002 (1991). Dactylaria manifesta R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 29 (1991). Dactylaria monticola R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 30 (1991). Dactylaria nectandrae R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 30 (1991). Dactylaria nervicola H. P. Upadhyay & Mankau, Mycologia 83, 371 (1991). Dactylaria queenslandica Matsush., Matsushima Mycological Memoirs 6, 15 (1989). Dactylaria quercicola W. B. Kendr. & R. F. Castan4 eda, University of Waterloo Biology Series 35, 31 (1991). Dactylaria sandinensis R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 32 (1991).
1268 Dactylaria simonensis R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 32 (1991). Dactylaria sparsa R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 33 (1991). Dactylaria splendida R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 33 (1991). Dactylaria sporexamorpha Glockling, Mycotaxon 43, 295 (1992). 3 Dactylaria pyriformis Tzean & J. L. Chen, Mycological Research 95, 1002 (1991), non Dactylaria pyriformis Juniper, Transactions of the British Mycological Society 37, 437 (1954). Dactylaria triseptata (Matsush.) R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 47 (1991). 3 Pleurophragmium triseptatum Matsush., Icones Microfungorum a Matsushima Lectorum, p. 116 (1975). Dactylaria tunicata Goh & K. D. Hyde, sp. nov. Dactylaria xinjiangensis Z. Jiao, X. Z. Liu & Y. T. Wang, Mycotaxon 52, 113 (1994). Dactylaria zapatensis R. F. Castan4 eda, Fungi Cubenses III, 5 (1988).
keys and illustrations for the 41 Dactylaria species he reviewed (Table 2), they are not reported here. TAXONOMY Dactylaria tunicata Goh & K. D. Hyde, sp. nov. (Figs 1–7, 10) Etym. : from Latin tunica meaning ‘ coating ’, referring to the characteristic hyaline gelatinous sheath of the conidia Coloniae effusae, griseolae vel olivaceae. Mycelium in ligno substrato partim superficiale et partim immersum, ex hyphis pallide vel modice brunneis, 2–3 µm diam., septatis, ramosis compositum. Stromata nulla. Setae et hyphopodia absunt. Conidiophora macronemata, solitaria vel laxe fasciculata, erecta, cylindrica, recta vel flexuosa, non ramosa, laevia, 2–6-septata, interdum ad septa leniter constricta, 75–160 µm longa, ad basim inflata et 8–10 µm lata, superne 4–4±5 µm lata, in latitudinibus uniformia vel apicem versus leniter attenuata, modice olivaceo-brunnea, apicem versus pallide olivacea vel subhyalina. Cellulae conidiogenae in conidiophoris incorporatae, terminales, cylindricae, subhyalinae, denticulatae. Conidia acropleurogena, holoblastica, solitaria, naviculata vel fusiformia, plerumque a latere parallelia, extrema acuminata, hyalina, uniseptata, 25–31¬3–4±5 µm, tunica gelatinosa hyalina fragili praedita ; conidiorum secessio schizolytica. Holotypus : Australia : north Queensland, Cowbay, in ligno putrido submerso in rivulo, April 1995, T. M. Hyde et K. D. Hyde (HKU(M) 2253).
Colonies effuse, greyish to olivaceous. Mycelium partly superficial and partly immersed in woody substratum, composed of pale to mid brown, 2–3 µm diam., septate, branched hyphae. Stromata none. Setae and hyphopodia absent. Conidiophores macronematous, solitary or 2–3 divergently fasciculate, erect, cylindrical, straight or flexuous, unbranched, smooth-walled, 2–6-septate, sometimes slightly constricted at the septa, 75–160 µm long, swollen and 8–10 µm wide at the base, stalk 4–4±5 µm wide, uniform in width or slightly attenuate towards the apex, mid olivaceous brown, becoming pale olivaceous to subhyaline towards the denticulate apex. Conidiogenous cells integrated, terminal, cylindrical, subhyaline, proliferating sympodially, with cylindrical denticles ; denticles hyaline, 1–2±5 µm long, ca 0±5– 1 µm wide. Conidial secession schizolytic. Conidia acropleurogenous, holoblastic, solitary, naviculate to fusiform, mostly with parallel sides, conically acuminate at both ends,
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Figs 8–27. Conidia of Dactylaria spp. Fig. 8. D. endophytica. Fig. 9. D. iriomoteana. Fig. 10. D. tunicata. Fig. 11. D. havanensis. Fig. 12. D. hemibeltranioidea. Fig. 13. D. sandinensis. Fig. 14. D. longidentata. Fig. 15. D. cubensis. Fig. 16. D. sporexamorpha. Fig. 17. D. appendiculata. Fig. 18. D. argentina. Fig. 19. D. attenuata. Fig. 20. D. quercicola. Fig. 21. D. botulispora. Fig. 22. D. fecundissima. Fig. 23. D. fabiformis. Fig. 24. D. lunata. Fig. 25. D. isoscelispora. Fig. 26. D. manifesta. Fig. 27. D. xinjiangensis. Scale bars, 5 µm.
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Figs 28–44. Conidia of Dactylaria spp. Fig. 28. D. simonensis. Fig. 29. D. hoogii. Fig. 30. D. lignicola. Fig. 31. D. eucalypti. Fig. 32. D. zapatensis. Fig. 33. D. splendida. Fig. 34. D. nectandrae. Fig. 35. D. cylindrospora. Fig. 36. D. triseptata. Fig. 37. D. queenslandica. Fig. 38. D. longispora. Fig. 39. D. sparsa. Fig. 40. D. leptosphaeriicola. Fig. 41. D. monticola. Fig. 42. D. aspirensis. Fig. 43. D. nervicola. Fig. 44. D. arecae. Scale bars, 5 µm.
hyaline, uniseptate, 25–31¬3–4±5 µm, winged with a hyaline, fragile, gelatinous sheath.
conidiophores with conspicuous, cylindrical denticles and hyaline conidia.
Note : This species could be classified under the Section Diplorhinotrichum (sensu de Hoog) as it produces dematiaceous
Other material examined : HKU (M) 2238, April 1995, T. M. Hyde and K. D. Hyde, on decaying wood submerged in freshwater stream, Cowbay, north Queensland, Australia.
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Key to Dactylaria species on plant material post de Hoog (1985) 1. As an endophyte in living leaves of Prunus (Rosaceae) ; conidiogenous loci flattened, 2–3±5 µm wide ; conidia cylindro-clavate, 0–1septate, septum located nearer the base, 6–17¬2–4±8 µm . . . . . . . . . . . D. endophytica 1. As a saprotroph on decaying plant parts or combination of characters not as above . . . . . . . . 2. 2. Two forms of conidia present : cylindro-ellipsoid conidia 2–4-septate, 18–30¬5–6 µm ; narrowly obclavate conidia 2–6-septate, 30–60¬5–6 µm ; conidial secession rhexolytic ; occurring on decaying leaves of Pandanus . . . . . D. iriomoteana 2. Only one form of conidia present or combination of characters not as above . . . . . . . . . 3. 3. Conidia with a distinct hyaline mucilaginous sheath, hyaline, 25–31¬3–4±5 µm, acerose, navicular to fusiform, uniseptate ; occurring on submerged wood . . . . . . . . . . . . . . . . . . D. tunicata 3. Conidia without mucilaginous sheath or combination of characters not as above . . . . . . . . . 4. 4. Conidia unicellular or rarely uniseptate . . . . . . . . . . . . . . . . 5. 4. Conidia predominantly septate . . . . . . . . . . . . . . . . . 8. 5. Conidia more or less cylindrical, naviculate to fusiform, with length to width ratio 6–8 : 1 ; occurring on decaying leaves of dicotyledons . . . . . . . . . . . . . . . . . . . . . 6. 5. Conidia more or less clavate to elliptical, with length to width ratio 3–4(–5) : 1 ; occurring on decaying wood or dead stems of grasses . . . . . . . . . . . . . . . . . . . . . . 7. 6. Conidia cylindro-clavate, inaequilateral, 9–11¬1–2 µm ; conidiophores hyaline, short, 9–12¬2–3 µm . . . D. havanensis 6. Conidia fusiform, cylindrical to naviculate, symmetrical, 12–20±5¬2–2±5 µm ; conidiophores brown, erect, 30–180¬4–8 µm . . . . . . . . . . . . . . . . . D. hemibeltranioidea 7. Conidia more or less clavate, apex papillate, 0-septate, eguttulate, 9–16¬3 µm . . . . . . . D. sandinensis 7. Conidia elliptical, apex apapillate, 0(–1)-septate, guttulate, 8–11¬2–3 µm . . . . . . . . D. longidentata 8. Conidia pyriform, uniseptate . . . . . . . . . . . . . . . . . . 9. 8. Conidia of other shapes . . . . . . . . . . . . . . . . . . . 10. 9. Conidiophores distinct, brown, 60–200¬3–4 µm ; conidia 19–25¬9–12 µm ; occurring on decaying leaves . . D. cubensis 9. Conidiophores reduced, hyaline, 7–22¬2±7–3±5 ; conidia 7±5–12±5¬3–5 µm ; occurring on decaying stems . D. sporexamorpha 10. Conidia 1–3-septate, obclavate to subcylindrical or fusiform, apex distinctly narrowed or attenuated, rostrate or appendiculate . . . . . . . . . . . . . . . . . . . . 11. 10. Conidia obclavate or of other shapes, apex not as above . . . . . . . . . . . . 14. 11. Conidia 4–5 µm wide, apex extended into a subulate or filiform appendage ; occurring on decaying wood submerged in freshwater habitat . . . . . . . . . . . . . . . . . . . . . . 12. 11. Conidia 2–3 µm wide, apex attenuated but not filiform ; occurring on dead leaves . . . . . . . . . 13. 12. Conidia subhyaline to pale brown, conidial body fusiform or subcylindrical, thick-walled, outer wall of apical cell extended into a filiform appendage ; conidiogenous denticles 1 µm wide . . . . . . . . . . D. appendiculata 12. Conidia hyaline, obclavate, thin-walled, apical cell attenuated into a subulate structure ; conidiogenous denticles 2–2±5 µm wide . . . . . . . . . . . . . . . . . . . D. argentina 13. Conidiophores hyaline, 1±5–2 µm wide ; conidia 1–2-septate, obclavate, attenuated smoothly towards the apex ; conidial secession rhexolytic . . . . . . . . . . . . . . . . . . . D. attenuata 13. Conidiophores brown, 2–3±5 µm wide ; conidia mostly 3-septate, more or less obclavate to ampulliform, constricted in the middle to form a tapered apical beak ; conidial secession schizolytic . . . . . . . . . . D. quercicola 14. Conidia distinctly curved, lunate to sublunate, allantoid, botuliform or fabiform, apex obtuse ; 1–2-septate . . . . 15. 14. Conidia more or less straight and of other shapes . . . . . . . . . . . . . 18. 15. Conidiophores brown, upper part ca 5 µm wide ; conidia sublunate or sausage-shaped . . . . . . . . 16. 15. Conidiophores hyaline, upper part ca 3 µm wide ; conidia allantoid or bean-shaped . . . . . . . . . 17. 16. Conidia botuliform, base obconic, 1-septate, 9–13¬2±5 µm . . . . . . . . . . D. botulispora 16. Conidia sublunate, base truncate, 2-septate, 11–25¬2±5–3 µm . . . . . . . . . D. fecundissima 17. Conidia 3±2–4 µm wide, ends rounded . . . . . . . . . . . . . . D. fabiformis 17. Conidia 2±3–3±3 µm wide, ends acute . . . . . . . . . . . . . . . D. luncata 18. Conidia narrowly cuneiform or elongated wedge-shaped, apex truncate or subtruncate and distinctly broader (3–3±5 µm wide), base 1–1±5 µm wide . . . . . . . . . . . . . . . . . . . 19. 18. Conidia of other shapes . . . . . . . . . . . . . . . . . . 20. 19. Conidia 1-septate . . . . . . . . . . . . . . . . . . D. isoscelispora 19. Conidia 2–5-septate . . . . . . . . . . . . . . . . . D. manifesta 20. Conidia predominantly with 4–5 or more septa . . . . . . . . . . . . . . 21. 20. Conidia uniseptate, 2–3-septate or rarely with up to 4 septa . . . . . . . . . . . . 24. 21. Conidia not exceeding 3±7 µm in width . . . . . . . . . . . . . . . . 22. 21. Conidia 4–5 µm wide . . . . . . . . . . . . . . . . . . . 23. 22. Apical portion of conidiophores irregularly broadened, denticles not protruding ; conidia fusiform, straight to slightly curved (3–)5(–7)-septate, 34±6–54±3¬2–3±7 µm ; habitat in orchard soil . . . . . . . . . D. xinjiangensis 22. Apical portion of conidiophores not broadened, denticles protruding ; conidia more or less cylindrical, curved, (3–)5(–8)-septate, 20–40¬2–2±5 µm ; habitat on dead stems of Cyperus . . . . . . . . . . . D. simonensis 23. Conidiogenous denticles prominent, 1–2 µm long ; conidia falcate or fusiform, apex acute, 4–8-septate, 30–45 µm long ; occurring on dead leaves and stems of Cyperus . . . . . . . . . . . . . . . D. hoogii 23. Conidiogenous denticles low in profile, broad, inconspicuous ; conidia fusiform, apex rounded, rounded, 5–9-septate, 60–65 µm long ; occurring on decaying wood . . . . . . . . . . . . . . . . D. lignicola 24. Conidia cylindrical to subcylindrical, with truncate or broadly rounded ends . . . . . . . . . 25. 24. Conidia navicular or fusiform to obclavate, with acute ends . . . . . . . . . . . . 33. 25. Conidia uniseptate to 2-septate . . . . . . . . . . . . . . . . . 26. 25. Conidia predominantly 3-septate . . . . . . . . . . . . . . . . . 29. 26. Conidiophores hyaline, reduced, 5–12¬1–2 µm ; conidia 1–2-septate, 10–26¬1–1±5 µm . . . . D. eucalypti 26. Conidiophores pale brown to brown, at least 12–100 µm long . . . . . . . . . . . 27.
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27. Conidiophoress up to 60 µm long, short, stalk 1–2 µm wide ; conidia 2-septate, 18–26¬1–1±5 µm . . . . D. zapatensis 27. Conidiophores up to 100 µm long, stalk at least 3 µm wide ; conidia 1-septate, 2–2±5 µm wide . . . . . . 28. 28. Conidiophores 5–8 µm wide ; conidia 13–16 µm long, subcylindrical or narrowly clavate, apex rounded, basal cell gently tapered and obconically truncate . . . . . . . . . . . . . . . . D. splendida 28. Conidiophores 3–4 µm wide ; conidia 16–20 µm long, almost perfectly cylindrical, apex rounded, basal cell cylindrical and truncate . . . . . . . . . . . . . . . . . . . D. nectandrae 29. Conidiophores robust, 100–220 µm long, 4–6±5 µm wide ; conidia subhyaline, pale brown to brown . . . . . 30. 29. Conidiophores up to 90 µm long, not exceeding 4 µm wide ; conidia hyaline . . . . . . . . . 31. . 30. Conidia pale brown to brown, versicoloured, end cells paler, 30–40¬5–6 µm . . . . . . . D. cylindrospora 30. Conidia subhyaline to mid olivaceous, uniform in colour, 14–25¬4–5 µm . . . . . . . D. triseptata 31. Conidiophores pale brown ; apex of conidia rounded and slightly swollen . . . . . . . . . . 32. 31. Conidiophores hyaline ; conidia 20–33¬2±8–4 µm, apex not swollen . . . . . . . . . D. queenslandica 32. Conidiophores 12–35¬3–4 µm ; conidia broader, 26–42¬2±5–4 µm, constricted at septa . . . . . D. longispora 32. Conidiophores 7–12¬3–4 µm ; conidia slender, 26–36¬1±5–2 µm, not constricted at septa . . . . . D. sparsa 33. Conidia predominately uniseptate, slender, not exceeding 2±5 µm in width . . . . . . . . . . 34. 33. Conidia predominantly 2–3-septate or rarely 4-septate, broader, 3–5±3 µm in width . . . . . . . . 35. 34. Conidiophores pale olivaceous to brown, 2±5–4 µm wide, conidia narrowly obclavate to fusiform, subhyaline to very pale olivaceous, 40–55¬1±5–2±5 µm ; occurring on pseudothecia of Leptosphaeria (Ascomycetes) . . . D. leptosphaeriicola 34. Conidiophores hyaline, 2–2±5 µm wide ; conidia narrowly fusiform, hyaline 30–37¬1–1±5 µm ; occurring on dead leaves . . . . . . . . . . . . . . . . . . . D. monticola 35. Apical portion of conidiophores attenuated ; conidiogenous cells terminal and intercalary, with sparse and scattered denticles (1–3 per cell) ; conidia naviculate to fusiform, 14–17¬3–4 µm . . . . . . . . . . . D. aspirensis 35. Apical portion of conidiophores broader with aggregated denticles or expanded into a denticulate vesicle ; conidia clavate, obclavate, ellipsoidal to fusiform, up to 4±5 µm or more in width . . . . . . . . . . . . . 36. 36. Apex of conidiophores enlarged into a denticulate vesicle ; denticles conical to tapered or sometimes inconspicuous ; conidia clavate to fusiform or ellipsoidal, 18±2–31±2¬3±8–5±3 µm ; occurring on decaying grasses (Poaceae) . . . D. nervicola 36. Apical denticulate vesicles lacking ; denticles cylindrical, conspicuous ; conidia obclavate to fusiform, 12–24¬3–4±5 µm ; occurring on decaying palm rachids (Arecaceae) . . . . . . . . . . . . . D. arecae
We are grateful to Dr Eric McKenzie (NZ) for the presubmission review of this work. Mr A. Y. P. Lee (HK) and Ms Helen Leung (HK) are thanked for their photographic and technical assistance. REFERENCES Arambarri, A., Cabello, M. & Mengascini, A. (1987). New hyphomycetes from Santiago River. II. (Buenos Aires Province, Argentina). Mycotaxon 30, 263–267. Bhatt, G. C. & Kendrick, W. B. (1968). The generic concept of Diplorhinotrichum and Dactylaria, and a new species of Dactylaria from soil. Canadian Journal of Botany 46, 1253–1257. Braun, U. & Crous, P. (1992). Dactylaria leptosphaeriicola spec. nov. Mycotaxon 45, 101–103. Castan4 eda Ruiz, R. F. (1988). Fungi Cubenses. III. Instituto de Investigaciones Fundamentales en Agricultura Tropica ‘ Alejandro de Humboldt ’ : Habana. Castan4 eda Ruiz, R. F. & Kendrick, W. B. (1990). Conidial fungi from Cuba. I. University of Waterloo Biological Series 32, 1–53. 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. Cazau, C., Arambarri, A. & Cabello, M. (1990). New hyphomycetes from Santiago River. IV. (Buenos Aires Province, Argentina). Mycotaxon 38, 21–25. Dixon, D. M. & Salkin, I. F. (1986). Morphologic and physiologic studies of three dematiaceous pathogens. Journal of Clinical Microbiology 24, 12–15. Dixon, D. M., Walsh, T. J., Salkin, I. F. & Polak, A. (1987). Dactylaria constricta : another dematiaceous fungus with neurotropic potential in mammals. Journal of Medical and Veterinary Mycology 25, 55–58. Glockling, S. L. (1992). The use of the binomial Dactylaria pyriformis. Mycotaxon 43, 295–296. Hoog, G. S. de (1985). Taxonomy of the Dactylaria-complex. IV. Dactylaria, Neta, Subulispora and Scolecobasidium. Studies in Mycology 26, 1–60. Hoog, G. S. de & Arx, J. A. von (1973). Revision of Scolecobasidium and Pleurophragmium. Kavaka 1, 55–60. Hoog, G. S. de & Hennebert, G. L. (1983). Taxonomy of the Dactylaria complex. III. A pleomorphic species of Isthmolongispora. Proceedings Koninklijke Nederlandse Akademie van Wetenschappen, Series C, 86, 343–346. (Accepted 28 January 1997)
Hoog, G. S. de & Oorschot, C. A. N. van (1985). Taxonomy of the Dactylariacomplex, VI. Key to the genera and check-list of epithets. Studies in Mycology 26, 97–122. Hoog, G. S. de, Oorschot, C. A. N. van & Hijwegen, T. (1983). Taxonomy of the Dactylaria-complex. II. Dissoconium gen. nov. and Cordana Preuss. Proceedings Koninklijke Nederlandse Akademie van Wetenschappen., Series C, 86, 197–206. Jiao, Z., Liu, X. Z. & Wang, Y. T. (1994). A new species of Dactylaria from China. Mycotaxon 52, 113–115. Matsushima, T. (1987). Matsushima Mycological Memoirs No. 5. Published by the author, Kobe. Matsushima, T. (1989). Matsushima Mycological Memoirs No. 6. Published by the author, Kobe. Matsushima, T. (1993). Matsushima Mycological Memoirs No. 7. Published by the author, Kobe. Matsushima, T. (1995). Matsushima Mycological Memoirs No. 8. Published by the author, Kobe. Oorschot, C. A. N. van (1985). Taxonomy of the Dactylaria-complex. V. A review of Arthrobotrys and allied genera. Studies in Mycology 26, 61–96. Rifai, M. A. (1968). The hyphomycete genus Dactylaria Sacc. Reinwardtia 7, 357–374. Saccardo, P. A. (1880). Conspectus generum fungorum Italiae inferiorum. Michelia 2, 1–38. Salkin, I. F. & Dixon, D. M. (1987). Dactylaria constricta : description of two varieties. Mycotaxon 29, 377–381. Schenck, S., Kendrick, W. B. & Pramer, D. (1977). A new nematode-trapping hyphomycete and a re-evaluation of Dactylaria and Arthrobotrys. Canadian Journal of Botany 55, 977–985. Tzean, S. S. & Chen, J. L. (1991). Two new species of Dactylaria from Taiwan. Mycological Research 95, 1000–1004. Upadhyay, H. P. & Mankau, R. (1991). Dactylaria nervicola sp. nov. and Exserohilum novae-zelandae comb. nov. from Mexico. Mycologia 83, 371–376. Vittal, B. P. R. & Dorai, M. (1991). Three new Deuteromycetes from Eucalyptus leaf litter. Mycological Research 95, 501–503. Wu, W., Sutton, B. C. & Gange, A. C. (1996). Dactylaria endophytica sp. nov., an endophyte from leaves of Prunus lusitanica. Mycological Research 100, 524–526.
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Printed in the United Kingdom
A revision of Dactylaria, with description of D. tunicata sp. nov. from submerged wood in Australia
T. K. G O H A N D K. D. H Y D E Department of Ecology and Biodiversity, The University of Hong Kong, Pokfulam Road, Hong Kong
Dactylaria tunicata sp. nov. from submerged wood in a freshwater stream from Australia is described and illustrated. It differs from all previously described species of Dactylaria in having uniseptate conidia with a hyaline sheath. The 41 species of Dactylaria accepted by de Hoog are listed, together with a further 41 validly published species of Dactylaria post de Hoog. A taxonomic key to 37 of the 41 post de Hoog species is provided, based on the literature, together with a composite illustration of their conidial morphology. The other four species are discussed, but they are not considered to be Dactylaria species (sensu de Hoog).
Since the establishment of Dactylaria Sacc. (1880), based on the lectotype species D. purpurella (Sacc.) Sacc., there have been numerous additions to the genus. Hyphomycetes which produce hyaline, thin-walled, 1–several septate conidia on sympodially proliferating, erect, hyaline or brown conidiophores, with more or less cylindrical denticles, are usually placed in the genus. These include lignicolous saprotroph, foliicolous plant pathogens, animal disease agents, fungicolous hyperparasites and nematophagous species. The generic concept has become confused, taxonomically and ecologically heterogeneous, though species distinction is somewhat easier. The heterogeneity of Dactylaria and similar fungi has provoked some early generic revision (Bhatt & Kendrick, 1968 ; Rifai, 1968 ; de Hoog & von Arx, 1973 ; Schenck, Kendrick & Pramer, 1977). Recent taxonomic revision of the Dactylaria-complex is given by de Hoog & Hennebert (1983), de Hoog, van Oorschot & Hijwegen (1983), de Hoog (1985), van Oorschot (1985), and de Hoog & van Oorschot (1985). In these publications, species disposed in Dactylaria (sensu lato) have been reidentified and reevaluated. The genus was redefined and subdivided into four sections (de Hoog, 1985), i.e. Dactylaria, Diplorhinotrichum (Ho$ hn.) de Hoog, Mirandina (G. Arnaud ex Matsush.) de Hoog, and Pleurophragmium (Costantin) de Hoog, with 37 allied genera, such as Arthrobotrys Corda, Duddingtonia R. C. Cooke, Geniculifera Rifai, Neta Shearer & J. L. Crane, Scolecobasidium E. V. Abbott, and Subulispora Tubaki. Fortyone species were accepted in Dactylaria (sensu de Hoog) and keys were given to groups of species in each section (de Hoog, 1985). Keys and an annotated check-list of published epithets were also provided to species of Neta, Subulispora and other allied genera of the Dactylaria-complex (de Hoog, 1985 ; de Hoog & van Oorschot, 1985). When grown in different culture media, many species of the
Dactylaria-complex exhibit morphological variations. In his review, de Hoog (1985) concluded that the taxonomy of these fungi is still in an exploratory phase and awaits a balanced understanding of their biology and variability. Although four sections were proposed to express the gross morphological differences within Dactylaria, de Hoog regarded the generic delimitation at present as artificial. Despite such generic revision by de Hoog, Matsushima (1987, 1989, 1993, 1995) still maintained Mirandina and Pleurophragmium as genera separate from Dactylaria (sensu de Hoog), describing several new species. De Hoog’s generic revision, with the acceptance of 41 species in Dactylaria (de Hoog, 1985 ; Table 1), represents a major contribution to fungal taxonomy. During the past decade, a further 40 species of Dactylaria (sensu de Hoog) have been validly published (Dixon & Salkin, 1986 ; Arambarri, Cabello & Mengascini, 1987 ; Dixon et al., 1987 ; Matsushima, 1987, 1989, 1993 ; Salkin & Dixon, 1987 ; Castan4 eda Ruiz, 1988 ; Castan4 eda Ruiz & Kendrick, 1990 ; Cazau, Arambarri & Cabello, 1990 ; Castan4 eda Ruiz & Kendrick, 1991 ; Tzean & Chen, 1991 ; Upadhyay & Mankau, 1991 ; Vittal & Dorai, 1991 ; Braun & Crous, 1992 ; Glockling, 1992 ; Jiao, Liu & Wang, 1994 ; Wu, Sutton & Gange, 1996). Most of the species are described from decaying plant material ; however, D. endophytica W. Wu, B. Sutton & Gange (1996) occurs as an endophyte from a living symptomless leaf of Prunus lusitanica. Dactylaria constricta (E. V. Abbott) D. M. Dixon & Salkin var. constricta, and the other variety D. constricta var. gallopava (Ivimey Cook) Salkin & D. M. Dixon (1987) are reported to be pathogens of birds and other animals, including man. Most species of Dactylaria (sensu lato) produce smooth, hyaline conidia which are usually fusiform, naviculate, or cylindrical in shape, but in some species they may be obpyriform, obclavate, fabiform, botuliform, or other shapes.
Revision of Dactylaria and one new species Table 1. Forty-one species of Dactylaria accepted by De Hoog Section Dactylaria Dactylaria acerosa Matsush., Icones Microfungorum a Matsushima Lectorum, p. 48 (1975). Dactylaria clavata Matsush., Icones Microfungorum a Matsushima Lectorum, p. 49 (1975). Dactylaria dimorphospora Veenb.-Rijks, Acta Botanica Neerlandica 22, 641 (1973). Dactylaria humicola G. C. Bhatt & W. B. Kendr. sensu Knauss & Alfieri, Proceedings of Florida States Horticultural Society 83, 441 (1970). 3 Dactylaria humicola G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). ¯ Dactylaria pseudoampulliformis Matsush., Icones Microfungorum a Matsushima Lectorum, p. 52 (1975). Dactylaria lanosa Malla & W. Gams, Persoonia 6, 193 (1971). Dactylaria ponapensis Matsush., Matsushima Mycological Memoirs 4, 7 (1985). Dactylaria purpurella (Sacc.) Sacc., Michelia 2, 20 (1880) ; Sylloge fungorum omnium hucusque cognitorum 4, 195 (1886). 3 Acrothecium purpurellum Sacc., Michelia 1, 75 (1877). Dactylaria scolecospora P. M. Kirk, Transactions of the British Mycological Society 81, 404 (1983). Section Diplorhinotrichum Dactylaria affinis (O. Rostr.) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). 3 Diplorhinotrichum affine O. Rostr., Dansk Botanisk Arkiv 2, 43 (1916). Dactylaria africana (S. Hughes) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). 3 Diplorhinotrichum africanum S. Hughes, Mycological Papers 47, 9 (1951). Dactylaria biseptata Matsush., Icones Microfungorum a Matsushima Lectorum, p. 48 (1975). Dactylaria candibula (Ho$ hn.) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1256 (1968). 3 Diplorhinotrichum candidulum Ho$ hn., Sitzungsberichte Akademie Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse Abteilung 111, 1040 (1902). Dactylaria chrysosperma (Sacc.) G. C. Bhatt & W. B. Kendr., Canadian Journal of Botany 46, 1257 (1968). 3 Rhinotrichum chrysospermum Sacc., Nuovo Giornale Botanico Italiano 8, 196 (1876). 3 Didymotrichum chrysospermum (Sacc.) Ho$ hn., Sitzungsberichte Akademie Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse Abteilung 123, 140 (1914). 3 Diplorhinotrichum chrysospermum (Sacc.) S. Hughes, Canadian Journal of Botany 36, 764 (1958). Dactylaria curviclavata Matsush., Matsushima Mycological Memoirs 3, 7 (1983). Dactylaria fulva R. Y. Roy & Gujarati, Lloydia 28, 53 (1965). ¯ Dactylaria indica Agarwal & Chowdhry, Current Science 46, 352 (1977). Dactylaria fusifera (Berk. & M. A. Curtis) de Hoog, Studies in Mycology 26, 17 (1985). 3 Rhinotrichum fusiferum Berk. & M. A. Curtis, Grevillea 3, 108 (1875). 3 Didymotrichum fusiferum (Berk. & M. A. Curtis) Linder, Lloydia 5, 207 (1942). Dactylaria inaequilatera Matsush., Matsushima Mycological Memoirs 3, 8 (1983). Dactylaria intermedia Matsush., Matsushima Mycological Memoirs 1, 25 (1980). Dactylaria naviculiformis Matsush., Icones Microfungorum a Matsushima Lectorum, p. 51 (1975). Dactylaria subuliphora Matsush., Icones Microfungorum a Matsushima Lectorum, p. 52 (1975). Dactylaria uniseptata Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 19 (1971).
1266 Section Pleurophragmium Dactylaria acerina Matsush., Matsushima Mycological Memoirs 3, 7 (1983). Dactylaria cymbiformis Matsush., Matsushima Mycological Memoirs 1, 25 (1980). Dactylaria dimorpha Matsush., Icones Microfungorum a Matsushima Lectorum, p. 49 (1975). Dactylaria dioscorae M. B. Ellis, More Dematiaceous Hyphomycetes, p. 171 (1976). Dactylaria fusarioidea Matsush., Icones Microfungorum a Matsushima Lectorum, p. 50 (1975). Dactylaria madrasensis Matsush., Matsushima Mycological Memoirs 3, 8 (1983). Dactylaria mitrata Matsush., Matsushima Mycological Memoirs 3, 8 (Sep. 1983). ¯ Hemibeltrania mitrata P. M. Kirk, Mycotaxon 18, 275 (Oct. 1983). Dactylaria mucronulata Ellis & Langl., Journal of Mycology 6, 35 (1890). Dactylaria kumamotoensis Matsush., Matsushima Mycological Memoirs 2, 5 (1981). Dactylaria parvispora (Preuss) de Hoog & Arx, Kavaka 1, 58 (1973). 3 Cordana parvispora Preuss, Linnaea 25, 728 (1852). 3 Pleurophragmium parvisporum (Preuss) Hol.-Jech., CeskaU Mykologie 36, 223 (1972). ¯ Pleurophragmium simplex (Berk. & Broome) S. Hughes, Canadian Journal of Botany 36, 798 (1958). 3 Acrothecium simplex Berk. & Broome, Annals and Magazine of Natural History III 7, 382 (1861). ¯ Acrothecium simplex Berk. & Broome var. elatum Grove, Journal of Botany, London 24, 203 (1886). ¯ Pleurophragmium bicolor Costantin, Les MuceU dineU eU s, p. 100 (1888). Section Mirandina Dactylaria breviphora Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 37 (1971). Dactylaria congregata de Hoog, Studies in Mycology 26, 30 (1985). Dactylaria fragilis de Hoog, Studies in Mycology 26, 30 (1985). Dactylaria fusiformis Shearer & J. L. Crane, Mycologia 63, 243 (March, 1971) ; non Dactylaria fusiformis (Matsush.) de Hoog & Arx, Kavaka 1, 58 (1973). ¯ Mirandina typica Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 38 (Sep. 1971). Dactylaria irregularis de Hoog, Studies in Mycology 26, 32 (1985). Dactylaria lepida Minter, Transactions of the British Mycological Society 74, 570 (1980). Dactylaria obtriangularia Matsush., Icones Microfungorum a Matsushima Lectorum, p. 51 (1975). Doubtful species Mirandina corticola G. Arnaud ex Matsush., Icones Microfungorum a Matsushima Lectorum, p. 96 (1975). 3 Mirandina corticola G. Arnaud, Bulletin de la SocieU teU Mycologique de France 68, 206 (1952) (nomen nudum sine descriptione latina : Article 36, ICBN). ¯ Mirandina yushanensis Matsush., Matsushima Mycological Memoirs 3, 12 (1983). The range of variability of this species is not certain. The shapes and sizes of the conidiophores, denticles and conidia were found to have slight differences based on materials on natural substrate and in culture. Mirandina cylindrospora Matsush., Microfungi of the Solomon Islands and Papua–New Guinea, p. 37 (1971). This species resembles D. biseptata and D. fulva, differing in having conidia with up to 3 septa. Mirandina dactylelloides Matsush., Matsushima Mycological Memoirs 1, 50 (1970). This species is closer to D. dimorphospora, but, it also resembles Trichoconis echinophilum (C. Massal.) de Hoog, Fusarium pallidoroseum (Cooke) Sacc. (Pseudohansfordia venezuelensis (Deighton & Piroz.) de Hoog), and species of Vermispora Deighton & Piroz.
T. K. Goh and K. D. Hyde
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Figs 1–5. Dactylaria tunicata. Figs 1–2. Squash mounts of conidiophores and conidia in lactophenol. Figs 3–5. Conidia. Note the hyaline gelatinous sheath around the conidia. Scale bars : 1, 50 µm ; 2, 20 µm ; 3–5, 10 µm.
Dactylaria appendiculata Cazau, Arambarri & Cabello (1990) and D. argentina (Aramb. & Mengasc.) Cabello & Cazau (Cazau et al., 1990) produce conidia with an appendiculate apex or filiform extension of the apical cell. Dactylaria isthmospora Cabello, Mengasc. & Aramb. (Arambarri et al., 1987) is somewhat atypical because its conidiophores bear 3–5 terminal and intercalary denticles which are arranged in whorls, and the conidia are somewhat clavate with a globose apex connected by an isthmus. Dactylaria laxa R. F. Castan4 eda (1988) is another atypical species which produces two-celled, fusiform, greyish, verrucose conidia from conidiphores which are pigmented, multiseptate and geniculate, with terminal and intercalary, chiefly monodenticulate conidiogenous cells. It would be more suitable to place this species in Scolecobasidium E. V. Abbott or Ochroconis de Hoog & Arx (1973). During our investigation of freshwater microfungi occurring on submerged woody material from streams in Queensland, Australia, we found a hyphomycete representing a member of the Dactylaria-complex which we consider to be new to science. In this paper, we describe this species as Dactylaria tunicata sp. nov. (Figs 1–7). It is unique amongst Dactylaria species in having navicular, hyaline, 1-septate conidia with a hyaline gelatinous sheath, and belongs to the section Diplorhinotrichum (Ho$ hn.) de Hoog (1985) as it produces pigmented conidiophores with conspicuous denticles at the apical region. With the description of this new hyphomycete in Dactylaria, there are presently 82 accepted species in the genus. A list of all 41 Dactylaria species which have been validly published post de Hoog (1985) is given in Table 2. In this paper, we do not attempt to reclassify each species according to the emended generic concept of Dactylaria (sensu de Hoog), although we realized that the genus is heterogeneous. A taxonomic key to 37 of those species growing on plant material (published post de Hoog, 1985) is given here
Figs 6–7. Dactylaria tunicata, diagramatic representation. Fig. 6. Conidiophores bearing conidia. Fig. 7. Conidia with gelatinous sheath. Scale bar, 10 µm.
based on the literature, and a composite illustration of their conidial morphology is provided (Figs 8–44). The atypical species D. isthmospora, D. laxa, as well as the two varieties of D. constricta (not foliicolous or lignicolous species) are excluded as we do not believe these are true Dactylaria species (sensu de Hoog). Since de Hoog (1985) has already provided
Revision of Dactylaria and one new species Table 2. Species of Dactylaria published post de Hoog, 1985 ; de Hoog & van Oorschot, 1985 Dactylaria appendiculata Cazau, Aramb. & Cabello, Mycotaxon 38, 21 (1990). Dactylaria arecae (Matsush.) R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 26 (1991). 3 Pleurophragmium arecae Matsush., Matsushima Mycological Memoirs 5, 23 (1987). Dactylaria argentina (Aramb. & Mengasc.) Cabello & Cazau, Mycotaxon 38, 22 (1990). 3 Subulispora argentina Aramb. & Mengasc., Mycotaxon 30, 264 (1987). Dactylaria aspirensis R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 26 (1991). Dactylaria attenuata Matsush., Matsushima Mycological Memoirs 5, 10 (1987). Dactylaria botulispora R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 32, 13 (1990). Dactylaria constricta (E. V. Abbott) D. M. Dixon & Salkin var. constricta, Mycotaxon 29, 379 (1987). 3 Scolecobasidium constrictum E. V. Abbott, Mycologia 19, 30 (1927). Dactylaria constricta var. gallopava (Ivimey Cook) Salkin & D. M. Dixon, Mycotaxon 29, 379 (1987). 3 Diplorhinotrichum gallopavum Ivimey Cook, Sabouraudia 3, 241 (1964). Dactylaria cubensis R. F. Castan4 eda, Fungi Cubensis III, 4 (1988). Dactylaria cylindrospora (Matsush.) R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 27 (1991). 3 Pleurophragmium cylindrosporum Matsush., Icones Microfungorum a Matsushima Lectorum, p. 115 (1975). Dactylaria endophytica W. Wu, B. Sutton & Gange, Mycological Research 100, 524 (1996). Dactylaria eucalypti Vittal & Dorai, Mycological Research 95, 501 (1991). Dactylaria fabiformis Matsush., Matsushima Mycological Memoirs 7, 48 (1993). Dactylaria fecundissima R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 27 (1991). Dactylaria havanensis R. F. Castan4 eda, Fungi Cubenses III, 4 (1988). Dactylaria hemibeltranioidea R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 27 (1991). Dactylaria hoogii R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 28 (1991). Dactylaria iriomoteana Matsush., Matsushima Mycological Memoirs 5, 10 (1987). Dactylaria isoscelispora W. B. Kendr. & R. F. Castan4 eda, University of Waterloo Biology Series 32, 16 (1990). Dactylaria isthmospora Cabello, Mengasc. & Aramb., Mycotaxon 30, 263 (1987). Dactylaria laxa R. F. Castan4 eda, Fungi Cubenses III, 5 (1988). Dactylaria leptosphaeriicola U. Braun & Crous, Mycotaxon 45, 101 (1992). Dactylaria lignicola W. B. Kendr. & R. F. Castan4 eda, University of Waterloo Biology Series 35, 29 (1991). Dactylaria longidentata Cazau, Aramb. & Cabello, Mycotaxon 38, 22 (1990). Dactylaria longispora Matsush., Matsushima Mycological Memoirs 5, 10 (1987). Dactylaria lunata Tzean & J. L. Chen, Mycological Research 95, 1002 (1991). Dactylaria manifesta R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 29 (1991). Dactylaria monticola R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 30 (1991). Dactylaria nectandrae R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 30 (1991). Dactylaria nervicola H. P. Upadhyay & Mankau, Mycologia 83, 371 (1991). Dactylaria queenslandica Matsush., Matsushima Mycological Memoirs 6, 15 (1989). Dactylaria quercicola W. B. Kendr. & R. F. Castan4 eda, University of Waterloo Biology Series 35, 31 (1991). Dactylaria sandinensis R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 32 (1991).
1268 Dactylaria simonensis R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 32 (1991). Dactylaria sparsa R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 33 (1991). Dactylaria splendida R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 33 (1991). Dactylaria sporexamorpha Glockling, Mycotaxon 43, 295 (1992). 3 Dactylaria pyriformis Tzean & J. L. Chen, Mycological Research 95, 1002 (1991), non Dactylaria pyriformis Juniper, Transactions of the British Mycological Society 37, 437 (1954). Dactylaria triseptata (Matsush.) R. F. Castan4 eda & W. B. Kendr., University of Waterloo Biology Series 35, 47 (1991). 3 Pleurophragmium triseptatum Matsush., Icones Microfungorum a Matsushima Lectorum, p. 116 (1975). Dactylaria tunicata Goh & K. D. Hyde, sp. nov. Dactylaria xinjiangensis Z. Jiao, X. Z. Liu & Y. T. Wang, Mycotaxon 52, 113 (1994). Dactylaria zapatensis R. F. Castan4 eda, Fungi Cubenses III, 5 (1988).
keys and illustrations for the 41 Dactylaria species he reviewed (Table 2), they are not reported here. TAXONOMY Dactylaria tunicata Goh & K. D. Hyde, sp. nov. (Figs 1–7, 10) Etym. : from Latin tunica meaning ‘ coating ’, referring to the characteristic hyaline gelatinous sheath of the conidia Coloniae effusae, griseolae vel olivaceae. Mycelium in ligno substrato partim superficiale et partim immersum, ex hyphis pallide vel modice brunneis, 2–3 µm diam., septatis, ramosis compositum. Stromata nulla. Setae et hyphopodia absunt. Conidiophora macronemata, solitaria vel laxe fasciculata, erecta, cylindrica, recta vel flexuosa, non ramosa, laevia, 2–6-septata, interdum ad septa leniter constricta, 75–160 µm longa, ad basim inflata et 8–10 µm lata, superne 4–4±5 µm lata, in latitudinibus uniformia vel apicem versus leniter attenuata, modice olivaceo-brunnea, apicem versus pallide olivacea vel subhyalina. Cellulae conidiogenae in conidiophoris incorporatae, terminales, cylindricae, subhyalinae, denticulatae. Conidia acropleurogena, holoblastica, solitaria, naviculata vel fusiformia, plerumque a latere parallelia, extrema acuminata, hyalina, uniseptata, 25–31¬3–4±5 µm, tunica gelatinosa hyalina fragili praedita ; conidiorum secessio schizolytica. Holotypus : Australia : north Queensland, Cowbay, in ligno putrido submerso in rivulo, April 1995, T. M. Hyde et K. D. Hyde (HKU(M) 2253).
Colonies effuse, greyish to olivaceous. Mycelium partly superficial and partly immersed in woody substratum, composed of pale to mid brown, 2–3 µm diam., septate, branched hyphae. Stromata none. Setae and hyphopodia absent. Conidiophores macronematous, solitary or 2–3 divergently fasciculate, erect, cylindrical, straight or flexuous, unbranched, smooth-walled, 2–6-septate, sometimes slightly constricted at the septa, 75–160 µm long, swollen and 8–10 µm wide at the base, stalk 4–4±5 µm wide, uniform in width or slightly attenuate towards the apex, mid olivaceous brown, becoming pale olivaceous to subhyaline towards the denticulate apex. Conidiogenous cells integrated, terminal, cylindrical, subhyaline, proliferating sympodially, with cylindrical denticles ; denticles hyaline, 1–2±5 µm long, ca 0±5– 1 µm wide. Conidial secession schizolytic. Conidia acropleurogenous, holoblastic, solitary, naviculate to fusiform, mostly with parallel sides, conically acuminate at both ends,
T. K. Goh and K. D. Hyde
1269
Figs 8–27. Conidia of Dactylaria spp. Fig. 8. D. endophytica. Fig. 9. D. iriomoteana. Fig. 10. D. tunicata. Fig. 11. D. havanensis. Fig. 12. D. hemibeltranioidea. Fig. 13. D. sandinensis. Fig. 14. D. longidentata. Fig. 15. D. cubensis. Fig. 16. D. sporexamorpha. Fig. 17. D. appendiculata. Fig. 18. D. argentina. Fig. 19. D. attenuata. Fig. 20. D. quercicola. Fig. 21. D. botulispora. Fig. 22. D. fecundissima. Fig. 23. D. fabiformis. Fig. 24. D. lunata. Fig. 25. D. isoscelispora. Fig. 26. D. manifesta. Fig. 27. D. xinjiangensis. Scale bars, 5 µm.
Revision of Dactylaria and one new species
1270
Figs 28–44. Conidia of Dactylaria spp. Fig. 28. D. simonensis. Fig. 29. D. hoogii. Fig. 30. D. lignicola. Fig. 31. D. eucalypti. Fig. 32. D. zapatensis. Fig. 33. D. splendida. Fig. 34. D. nectandrae. Fig. 35. D. cylindrospora. Fig. 36. D. triseptata. Fig. 37. D. queenslandica. Fig. 38. D. longispora. Fig. 39. D. sparsa. Fig. 40. D. leptosphaeriicola. Fig. 41. D. monticola. Fig. 42. D. aspirensis. Fig. 43. D. nervicola. Fig. 44. D. arecae. Scale bars, 5 µm.
hyaline, uniseptate, 25–31¬3–4±5 µm, winged with a hyaline, fragile, gelatinous sheath.
conidiophores with conspicuous, cylindrical denticles and hyaline conidia.
Note : This species could be classified under the Section Diplorhinotrichum (sensu de Hoog) as it produces dematiaceous
Other material examined : HKU (M) 2238, April 1995, T. M. Hyde and K. D. Hyde, on decaying wood submerged in freshwater stream, Cowbay, north Queensland, Australia.
T. K. Goh and K. D. Hyde
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Key to Dactylaria species on plant material post de Hoog (1985) 1. As an endophyte in living leaves of Prunus (Rosaceae) ; conidiogenous loci flattened, 2–3±5 µm wide ; conidia cylindro-clavate, 0–1septate, septum located nearer the base, 6–17¬2–4±8 µm . . . . . . . . . . . D. endophytica 1. As a saprotroph on decaying plant parts or combination of characters not as above . . . . . . . . 2. 2. Two forms of conidia present : cylindro-ellipsoid conidia 2–4-septate, 18–30¬5–6 µm ; narrowly obclavate conidia 2–6-septate, 30–60¬5–6 µm ; conidial secession rhexolytic ; occurring on decaying leaves of Pandanus . . . . . D. iriomoteana 2. Only one form of conidia present or combination of characters not as above . . . . . . . . . 3. 3. Conidia with a distinct hyaline mucilaginous sheath, hyaline, 25–31¬3–4±5 µm, acerose, navicular to fusiform, uniseptate ; occurring on submerged wood . . . . . . . . . . . . . . . . . . D. tunicata 3. Conidia without mucilaginous sheath or combination of characters not as above . . . . . . . . . 4. 4. Conidia unicellular or rarely uniseptate . . . . . . . . . . . . . . . . 5. 4. Conidia predominantly septate . . . . . . . . . . . . . . . . . 8. 5. Conidia more or less cylindrical, naviculate to fusiform, with length to width ratio 6–8 : 1 ; occurring on decaying leaves of dicotyledons . . . . . . . . . . . . . . . . . . . . . 6. 5. Conidia more or less clavate to elliptical, with length to width ratio 3–4(–5) : 1 ; occurring on decaying wood or dead stems of grasses . . . . . . . . . . . . . . . . . . . . . . 7. 6. Conidia cylindro-clavate, inaequilateral, 9–11¬1–2 µm ; conidiophores hyaline, short, 9–12¬2–3 µm . . . D. havanensis 6. Conidia fusiform, cylindrical to naviculate, symmetrical, 12–20±5¬2–2±5 µm ; conidiophores brown, erect, 30–180¬4–8 µm . . . . . . . . . . . . . . . . . D. hemibeltranioidea 7. Conidia more or less clavate, apex papillate, 0-septate, eguttulate, 9–16¬3 µm . . . . . . . D. sandinensis 7. Conidia elliptical, apex apapillate, 0(–1)-septate, guttulate, 8–11¬2–3 µm . . . . . . . . D. longidentata 8. Conidia pyriform, uniseptate . . . . . . . . . . . . . . . . . . 9. 8. Conidia of other shapes . . . . . . . . . . . . . . . . . . . 10. 9. Conidiophores distinct, brown, 60–200¬3–4 µm ; conidia 19–25¬9–12 µm ; occurring on decaying leaves . . D. cubensis 9. Conidiophores reduced, hyaline, 7–22¬2±7–3±5 ; conidia 7±5–12±5¬3–5 µm ; occurring on decaying stems . D. sporexamorpha 10. Conidia 1–3-septate, obclavate to subcylindrical or fusiform, apex distinctly narrowed or attenuated, rostrate or appendiculate . . . . . . . . . . . . . . . . . . . . 11. 10. Conidia obclavate or of other shapes, apex not as above . . . . . . . . . . . . 14. 11. Conidia 4–5 µm wide, apex extended into a subulate or filiform appendage ; occurring on decaying wood submerged in freshwater habitat . . . . . . . . . . . . . . . . . . . . . . 12. 11. Conidia 2–3 µm wide, apex attenuated but not filiform ; occurring on dead leaves . . . . . . . . . 13. 12. Conidia subhyaline to pale brown, conidial body fusiform or subcylindrical, thick-walled, outer wall of apical cell extended into a filiform appendage ; conidiogenous denticles 1 µm wide . . . . . . . . . . D. appendiculata 12. Conidia hyaline, obclavate, thin-walled, apical cell attenuated into a subulate structure ; conidiogenous denticles 2–2±5 µm wide . . . . . . . . . . . . . . . . . . . D. argentina 13. Conidiophores hyaline, 1±5–2 µm wide ; conidia 1–2-septate, obclavate, attenuated smoothly towards the apex ; conidial secession rhexolytic . . . . . . . . . . . . . . . . . . . D. attenuata 13. Conidiophores brown, 2–3±5 µm wide ; conidia mostly 3-septate, more or less obclavate to ampulliform, constricted in the middle to form a tapered apical beak ; conidial secession schizolytic . . . . . . . . . . D. quercicola 14. Conidia distinctly curved, lunate to sublunate, allantoid, botuliform or fabiform, apex obtuse ; 1–2-septate . . . . 15. 14. Conidia more or less straight and of other shapes . . . . . . . . . . . . . 18. 15. Conidiophores brown, upper part ca 5 µm wide ; conidia sublunate or sausage-shaped . . . . . . . . 16. 15. Conidiophores hyaline, upper part ca 3 µm wide ; conidia allantoid or bean-shaped . . . . . . . . . 17. 16. Conidia botuliform, base obconic, 1-septate, 9–13¬2±5 µm . . . . . . . . . . D. botulispora 16. Conidia sublunate, base truncate, 2-septate, 11–25¬2±5–3 µm . . . . . . . . . D. fecundissima 17. Conidia 3±2–4 µm wide, ends rounded . . . . . . . . . . . . . . D. fabiformis 17. Conidia 2±3–3±3 µm wide, ends acute . . . . . . . . . . . . . . . D. luncata 18. Conidia narrowly cuneiform or elongated wedge-shaped, apex truncate or subtruncate and distinctly broader (3–3±5 µm wide), base 1–1±5 µm wide . . . . . . . . . . . . . . . . . . . 19. 18. Conidia of other shapes . . . . . . . . . . . . . . . . . . 20. 19. Conidia 1-septate . . . . . . . . . . . . . . . . . . D. isoscelispora 19. Conidia 2–5-septate . . . . . . . . . . . . . . . . . D. manifesta 20. Conidia predominantly with 4–5 or more septa . . . . . . . . . . . . . . 21. 20. Conidia uniseptate, 2–3-septate or rarely with up to 4 septa . . . . . . . . . . . . 24. 21. Conidia not exceeding 3±7 µm in width . . . . . . . . . . . . . . . . 22. 21. Conidia 4–5 µm wide . . . . . . . . . . . . . . . . . . . 23. 22. Apical portion of conidiophores irregularly broadened, denticles not protruding ; conidia fusiform, straight to slightly curved (3–)5(–7)-septate, 34±6–54±3¬2–3±7 µm ; habitat in orchard soil . . . . . . . . . D. xinjiangensis 22. Apical portion of conidiophores not broadened, denticles protruding ; conidia more or less cylindrical, curved, (3–)5(–8)-septate, 20–40¬2–2±5 µm ; habitat on dead stems of Cyperus . . . . . . . . . . . D. simonensis 23. Conidiogenous denticles prominent, 1–2 µm long ; conidia falcate or fusiform, apex acute, 4–8-septate, 30–45 µm long ; occurring on dead leaves and stems of Cyperus . . . . . . . . . . . . . . . D. hoogii 23. Conidiogenous denticles low in profile, broad, inconspicuous ; conidia fusiform, apex rounded, rounded, 5–9-septate, 60–65 µm long ; occurring on decaying wood . . . . . . . . . . . . . . . . D. lignicola 24. Conidia cylindrical to subcylindrical, with truncate or broadly rounded ends . . . . . . . . . 25. 24. Conidia navicular or fusiform to obclavate, with acute ends . . . . . . . . . . . . 33. 25. Conidia uniseptate to 2-septate . . . . . . . . . . . . . . . . . 26. 25. Conidia predominantly 3-septate . . . . . . . . . . . . . . . . . 29. 26. Conidiophores hyaline, reduced, 5–12¬1–2 µm ; conidia 1–2-septate, 10–26¬1–1±5 µm . . . . D. eucalypti 26. Conidiophores pale brown to brown, at least 12–100 µm long . . . . . . . . . . . 27.
Revision of Dactylaria and one new species
1272
27. Conidiophoress up to 60 µm long, short, stalk 1–2 µm wide ; conidia 2-septate, 18–26¬1–1±5 µm . . . . D. zapatensis 27. Conidiophores up to 100 µm long, stalk at least 3 µm wide ; conidia 1-septate, 2–2±5 µm wide . . . . . . 28. 28. Conidiophores 5–8 µm wide ; conidia 13–16 µm long, subcylindrical or narrowly clavate, apex rounded, basal cell gently tapered and obconically truncate . . . . . . . . . . . . . . . . D. splendida 28. Conidiophores 3–4 µm wide ; conidia 16–20 µm long, almost perfectly cylindrical, apex rounded, basal cell cylindrical and truncate . . . . . . . . . . . . . . . . . . . D. nectandrae 29. Conidiophores robust, 100–220 µm long, 4–6±5 µm wide ; conidia subhyaline, pale brown to brown . . . . . 30. 29. Conidiophores up to 90 µm long, not exceeding 4 µm wide ; conidia hyaline . . . . . . . . . 31. . 30. Conidia pale brown to brown, versicoloured, end cells paler, 30–40¬5–6 µm . . . . . . . D. cylindrospora 30. Conidia subhyaline to mid olivaceous, uniform in colour, 14–25¬4–5 µm . . . . . . . D. triseptata 31. Conidiophores pale brown ; apex of conidia rounded and slightly swollen . . . . . . . . . . 32. 31. Conidiophores hyaline ; conidia 20–33¬2±8–4 µm, apex not swollen . . . . . . . . . D. queenslandica 32. Conidiophores 12–35¬3–4 µm ; conidia broader, 26–42¬2±5–4 µm, constricted at septa . . . . . D. longispora 32. Conidiophores 7–12¬3–4 µm ; conidia slender, 26–36¬1±5–2 µm, not constricted at septa . . . . . D. sparsa 33. Conidia predominately uniseptate, slender, not exceeding 2±5 µm in width . . . . . . . . . . 34. 33. Conidia predominantly 2–3-septate or rarely 4-septate, broader, 3–5±3 µm in width . . . . . . . . 35. 34. Conidiophores pale olivaceous to brown, 2±5–4 µm wide, conidia narrowly obclavate to fusiform, subhyaline to very pale olivaceous, 40–55¬1±5–2±5 µm ; occurring on pseudothecia of Leptosphaeria (Ascomycetes) . . . D. leptosphaeriicola 34. Conidiophores hyaline, 2–2±5 µm wide ; conidia narrowly fusiform, hyaline 30–37¬1–1±5 µm ; occurring on dead leaves . . . . . . . . . . . . . . . . . . . D. monticola 35. Apical portion of conidiophores attenuated ; conidiogenous cells terminal and intercalary, with sparse and scattered denticles (1–3 per cell) ; conidia naviculate to fusiform, 14–17¬3–4 µm . . . . . . . . . . . D. aspirensis 35. Apical portion of conidiophores broader with aggregated denticles or expanded into a denticulate vesicle ; conidia clavate, obclavate, ellipsoidal to fusiform, up to 4±5 µm or more in width . . . . . . . . . . . . . 36. 36. Apex of conidiophores enlarged into a denticulate vesicle ; denticles conical to tapered or sometimes inconspicuous ; conidia clavate to fusiform or ellipsoidal, 18±2–31±2¬3±8–5±3 µm ; occurring on decaying grasses (Poaceae) . . . D. nervicola 36. Apical denticulate vesicles lacking ; denticles cylindrical, conspicuous ; conidia obclavate to fusiform, 12–24¬3–4±5 µm ; occurring on decaying palm rachids (Arecaceae) . . . . . . . . . . . . . D. arecae
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