Mycal. Res. 97 (II): 1328-1332 (1993)
1328
Prinled in Greal Brilain
The genus Phyllachora from Australia: P. queenslandica and notes on P. apiculata from Neolitsea
CERIDWEN A. PEARCE AND KEVIN D. HYDE'" Division of Plant Protection, Queensland Department of Primary Industries, Mareeba, Qld 4880, Australia
Phyllachora queenslandica on Neolitsea dealbata is redescribed from north Queensland material and illustrated with differential interference micrographs. The status of Phyllachora apiculata from Neolitsea is also discussed.
Phyllachora queenslandica Hansf. (Hansford, 1956) was described from leaf specimens of Neolitsea dealbata (R. Br.) Merr., collected on Mt Buderim in Queensland, exhibiting brown lesions supporting shiny black tar spots. A similar tar spot on leaves of the same plant was collected in the rainforests around Lake Eacham on the Atherton Tablelands in north Queensland, and subsequent examination showed this collection to match Hansford's description of P. queenslandica. The material was, therefore, compared with the type specimen deposited by Hansford, and although this is in poor condition we consider the collections to represent the same taxon. The species is redescribed and illustrated with differential interference contrast (DIC) micrographs taken from our specimen. In a search of the literature for Phyllachora species on Neolitsea in Australia we found a further record; Phyllachora apiculata SacCo & Berl. (Simmonds, 1966). The fungus is based on Dothidella apiculata Sacco & Berl. (Saccardo & Berlese, 1885), which was listed as occurring in Queensland by Bailey (1886). It was later listed as Phyllachora apiculata SacCo & Ber!. by Bailey (1909), and although we can find no record of its synonymy with Phyllachora, it is the same record (Simmonds, 1966). The description of Dothidella apiculata given in Saccardo & Berlese (1885) gives the asci as being 4-spored and the ascospores bicelled, and it seems unlikely that this species could be a Phyllachora. We have examined material of D. apiculata (Australia: Queensland, Samford, on senescent leaves of Neolitsea dealbata, September 1883, Rev. B. Scortechini, PAD! holotype) and confirm this is not a Phyllachora. The asci are 8-spored, bitunicate, with a distinct apical ring; ascospores are hyaline, bicelled and with an eccentric septum, and ascomata form beneath a black, shiny, dome-shaped stroma. Its retention in Dothidella Speg., therefore, seems warranted
• Corresponding author, current address: Department of Botany, University of Hong Kong, Pokfulam Road, Hong Kong.
(Muller & Arx, 1962). The epithet 'apiculata' had also previously been used for P. apiculata Speg. from Panicum (Spegazzini, 1899), and the name used by Bailey (1909) therefore is illegitimate.
TAXONOMY Phyllachora queensIandica Hansf. Proc. Linn. Soc. N.S. W. 81,28 (1956). (Figs 1-19)
Leaf spot 1'4-2'3 mm diam., frequently coalescing, usually roughly circular, yellowish-brown, composed of an outer ring of degenerated host tissue with an inner area of roughly circular tar spots, 0'65-1'3 mm diam., solitary or gregarious, occasionally coalescing, shallowly domed, shiny to matt black, and sometimes forming a scattered ring. Leaf spot development most prominent on the upper leaf surface, the larger leaf spots often with a central region of effete ascomata (Figs 1, 2). Anamorph. A brown-black amorphous region, 40-50 ~m diam. composed of melanized host epidermal and palisade parenchyma cells and fungal hyphae, often forms on both sides of the leaf, with the conidiomata developing within this, immersed in the host mesophyll, often adjacent to ascomata (Fig. 5). Conidiomata 248-280 ~m diam., 170-288 ~m high, roughly globose, with a conical neck (Fig. 10). Conidiomatal wall distinct from teleomorph wall. composed of dark-brown to black flattened fungal cells and compressed host tissue. Upper wall to 34 ~m thick, merging in the palisade parenchyma with the blackened region above. Lower wall thinner, up to 20 ~m thick (Fig. I I). Fertile layer not clearly differentiated from the conidioma wall. Ostiole central and circular. Conidiogenous cells extend inwards from the base and sides of the conidiomatal wall, cylindrical, up to 2 ~m diam. at base, tapering to I ~m at the apex, 16-32 ~ long, hyaline, with conidia borne singly (Figs 7, 12). Conidia 5-6 x 1'8-2 ~m (i = 5'4 x 1'9 ~m, n = 20), hyaline, basilar, thin-walled and smooth, fractionally wider at the base, narrowing slightly to a rounded apex (Figs 8, 9).
Ceridwen A. Pearce and K. D. Hyde
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Figs 1-6. Differential interference contrast micrographs (except Figs 1,2) of Phyllachora queenslandica, teleomorph. Figs 1, 2. Tar spots. Figs 3, 5. Vertical sections through ascomata. The dose proximity of the anamorph is illustrated in Fig. 5. Fig. 4. Peridium. Fig. 6. Ascomatal neck. Scale bars: 1, 2 mm; 2, 1 mm; 3,5, 300 ~; 4, 6, 10 ~m.
Teleomorph. Ascomata 520-1625 ~ diam., 455-1275 ~m high (i = 963 X 863~, n = 20), immersed in the host mesophylL subglobose to ellipsoidal, base flattened, tapering apically to a conical neck (Figs 3, 5). Upper ascomatal wall to 10 ~m thick, composed of light-brown, thin-walled, flattened fungal cells and compressed host tissue (Fig. 4). Lower wall slightly thinner and similarly composed. A large, distinct, 85
dark-brown to black amorphous layer, comprised of melanized host epidermal and palisade cells and fungal hyphae, forms beneath the host cuticle. The ostiole is central, indistinct, and lined with fine, hyaline periphyses (Fig. 6). Asci and paraphyses arise from the base and lateral walls of the ascoma (Fig. 3). Paraphyses numerous, slightly longer than asci, not constricted at septa, thin-walled, branching, tapering from 4 ~ diam. at MYC 97
The genus Phyllachora from Australia
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Figs 7-12. Differential interference micrographs Phyllachora queenslandica, anamorph. Figs 7, 12. Conidiogenolls cells bearing conidia. Figs 8,9. Conidia. Fig. 10. Vertical section of conidioma. Fig. 11. Wall of conidioma. Scale bars: 7-9, 12, 5 11m; 10, 50 11m; 11, 25 J.1m.
base to 2 J.lm apically, with a rounded tip (Fig. 17). Asci 125-240 x 26-38 J.lm (i = 189 x 30 J.lm, n = 25), 8-spored, unitunicate, clavate, usually short-stalked, tapering to a rounded apex. No apical structure visible on staining (Figs 18,19). Ascospores 19-32 x 8-12'5 J.lm (i = 26 x 10'5 J.lm, n = 50), arranged loosely in ascus, uniseriate or biseriate, oblong-ellipticaL sometimes very slightly inequilateraL hyaline, thick-walled, guttulate, aseptate, encircled by a layer of mUcilage up to 16 J.lm thick (Figs 13-16). Host species: Neolitsea dealbata. Known distribution: Australia. Material examined: Australia: Queensland, Mt Buderim, on living leaves of Neolitsea dealbata, 1912, C. T. White, K, holotype, BRIP 3261, isotype; Atherton Tablelands, Lake Eacham area, on living leaves of Neolitsea dealbata, 15 Feb. 1992, C. A. Pearce, KDH 1033, BRIP 20460.
DISCUSSION Hansford (1956) did not report the presence of an anamorph in Phyllachora queenslandica, although one was observed in our collection. Hawksworth (1981) found that most genera of conidial fungi restricted to the Sphaeriales (then regarded as a group containing most dark-walled pyrenomycetes) were in fact parasites of the Phyllachorceae. Mycoparasitism of the Phyllachoraceae is common (Parbery, 1978; Sutton, 1964).
Cannon (1991) felt that this is possibly due to the longevity of the fruiting structures and their biotrophic existence. Saccardo (1883, 1928), Theissen & Sydow (1915), Doidge (1942), Orton (1944) and Cannon (1991) have each described species of Phyllachora with imperfect states being accepted as part of the same fungus due to the close proximity of the ascal and conidial fructifications. In P. queenslandica the mature conidiomata exist in close proximity to the immature teleomorph, the ascomata and conidiomata often developing beneath the same clypeus. For this reason we have accepted this anamorphic state as part of the P. queenslandica life cycle. In one instance we observed a mature conidioma release conidia through an ostiole in the lower leaf surface, while an adjacent mature ascoma released ascospores via its ostiole to the upper leaf surface. Cannon (1991) states that in many instances Phyllachora conidiomata may mature before the ascomata and continue to produce conidia for an extended period of time. The fresh specimen of P. queenslandica examined showed ascospores enclosed in a clear mucilaginous sheath up to 16 J.lm thick. It is particularly evident in water mounts at low magnification (e.g. x 100) (Fig. 13), or on staining with 1 % nigrosin (Figs. 14-16). The presence of mUCilaginous sheaths associated with ascospores is not uncommon in Phyllachora, although they may be ephemeral in nature and often difficult to detect in dried herbarium specimens (Cannon, 1991; Hyde
Ceridwen A. Pearce and K. D. Hyde
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Figs 13-19. Differential interference micrographs of Phyllachora quernslandica, teleomorph. Figs 13-16. Ascospores. Note the mucilaginous sheaths. Fig. 17. Squash illustrating asci and paraphyses. Figs 18,19. Asci which lack an apical ring. The mucilaginous sheath around the ascospores can be seen within the ascus. Scale bars: 13, 50 ~m; 14-19, 10 ~m.
& Cannon, 1992). Hansford (1956) notes in his original
description of the fungus that 'a mucilage envelope exists around the ascospores, up to 5-6 IJIIl thick'.
We thank the Northern Australian Quarantine Strategy and Australian Quarantine and Inspection Service for funding this research. Thanks also to the curators of K, BRIP and S for loan of the fungal material used in this study. John Clarkson is also thanked for identifying botanical specimens. John Alcorn is thanked for comments on the manuscript.
REFERENCES Bailey, F. M. (1886). Queensland Flora. A Synopsis of the First supplement. Government Printer: Brisbane, Qld, Australia. Bailey, F. M. (1909). Comprehensive Catalogue of Queensland Plants. Government Printer: Brisbane. Qld, Australia. Cannon, P. F. (1991). A revision of Phyllachora and some similar genera on the host family Leguminosae. Mycological Papers 163, 1-302. Doidge. E. M. (1942). Revised descriptions of South African species of Phyllachora and related genera. Bothalia 4, 421-463. Hansford, C. G. (1956). Australian fungi. III. Proceedings of the Linnean Society of New South Wales 81, 23-51. Hawksworth, D. L. (1981). A survey of the fungicolous conidia. In Biology of 8S-2
The genus Phyllachora from Australia Conidial Fungi (ed. G. T. Cole & W. B. Kendrick), pp. 171-244. Academic Press: London. Hyde, K. D. & Cannon, P. F. (1992). Polystigma sonneratiae causing leaf spots on the mangrove genus Sonneralia. Australian Systematic Botany S, 415-420. MUller, E. & Arx, J. (1962). Die Gattungen der didymosporen Pyrenomyceten. Beitriige zur Kryptogamenflora der Schweiz 11, 1-922. Orton, C. R. (1944). Graminicolous species of Phyllachora in North America. Mycologia 36, 18-53. Parbery, D. G. (1978). Phyllachora, Linochora and hyperparasites. In Taxonomy of Fungi (ed. C. V. Subramanian), pp. 263-277. University of Madras: Madras. Saccardo, P. A. (1883). Sylloge Fungorum. Vol. 2. Johnson Reprint Corporation: New York.
1332 Saccardo, P. A. (1928). Sylloge Fungorum. Vol. 24. Johnson Reprint Corporation: New York. Saccardo, P. A. & Berlese, A. N. (1885). Fungi Australienses. Revue Mycologique 7,92-98.
Simmonds, J. H. (1966). Host Index of Plant Diseases in Queensland. Queensland Department of Primary Industries: Brisbane, Australia. Spegazzini, C. (1899). Fungi Argentini novi v. critid. Linnaeus Press: Amsterdam. Sutton, B. C. (1964). Coelomycetes. III. Annellolacinia gen. nov., Aristastoma, Phaeocytostroma, Seimatosporium, etc. Mycological Papers 97, 1-42. Theissen, F. & Sydow, H. (1915). Die Dothideales. Kritisch-systematische Originaluntersuchungen. Annales Mycologici 13, 147-746.
(Accepted 5 April 1993)
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