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Johnston, P.R. 1999: Some hyperparasites of Meliolina in New Zealand. New Zealand Journal of Botany 37(2): 289-295.

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Johnston, P.R. 1999: Some hyperparasites of Meliolina in New Zealand. New Zealand Journal of Botany 37(2): 289-295.
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ADITIONAL SPECIMENS EXAMINED: NEW ZEALAND: SOUTH ISLAND: SOUTHLAND: Catlins, Lake Wilkie, on Meliolina leptospermi on Leptospermum scoparium, P. R. Johnston & A. Howarth, 9 May 1995, PDD 68411. STEWART ISLAND: Mason Bay, near start of track to Freshwater Landing, P. R. Johnston & E. H. C. McKenzie, 23 Nov 1998, PDD 69824.
DESCRIPTION: Conidiophores, often aggregated into white sporodochial-like masses, develop amongst the hyphae of Meliolina leptospermi colonies. The aerial hyphae of infected Meliolina colonies are sparse and eventually completely lost, the chlorotic host leaf tissue beneath the Meliolina colonies becoming exposed. The chlorotic area is speckled with remains of dark Meliolina hyphae within substomatal chambers. Mycelium 2.5-3 µm diam., thin-walled, pale brown, with short cells, irregularly convoluted and tangled, always amongst Meliolina hyphae but not closely adhering to them. Wine-red pigment diffuses into KOH from the mycelium. Conidiogenous cells held on the mycelium in irregular, compact groups. Conidiogenous cells 11-19 x 2.5-4 µm, hyaline, more or less cylindric, although narrower near apex, proliferation percurrent, with several rows of indistinct annellations. Conidia 44-48 x 3-3.5 µm, more or less fusoid, tapering to a truncate base, and attenuated into an elongate, flexuous apex, 3-septate, hyaline.
DIAGNOSIS: Conidiophora aggregata facientes sporodochia inter hyphas Meliolinas leptosperma; cellulae conidiogenae 11-19 x 2.5-4 µm; conidia 44-48 x 3-3.5 µm, fusiformes, 3-septata, basi truncatae, apici protracto flexuoso.

ETYMOLOGY: Refers to the genus of the host fungus.

NOTES: Annellospermosporella meliolinae, associated with sparse, partly eroded Meliolina leptospermi colonies, is often difficult to recognise macroscopically.
In two of the collections (PDD 63407, 69824), this species was often associated with perithecia of Malacaria meliolinae. In these collections, the mycelium from both species appears to exude a wine red pigment in KOH. In the third collection of A. meliolinae (PDD 68411), where Malacaria was not observed, the same KOH reaction occurs. Healthy Meliolina mycelium gives no reaction in KOH. Although the evidence is not strong, there is a possibility that these two species represent the anamorph and teleomorph of the same fungus. Hansford (1946) speculated that the anamorph ofMalacaria flagellata (Hansf.) Hansf. (M. meholicola Syd., see Rossman 1987) may be Atractalina parasitica (G.Winter) Deighton & Piroz. (as Arthrobotryum parasiticum (G.Winter) Hansf), primarily on the basis of the fungi being found in close proximity on the same host fungus colonies.

HOLOTYPUS: New Zealand, Auckland, North Shore City, Torbay, Stredwick Drive Reserve, on Meliolina leptospermi on Leptospermum scoparium, P. R. Johnston, 14 Sep 1994, PDD 63407.
DIAGNOSIS: Ab Spermosporella cellulis conidiogenis percurrentibus annellidis differens.

ETYMOLOGY: Similar to Spermosporella but with annellidic conidiogenous cells.

NOTES: Annellospermosporella is biologically and morphologically very similar to Spermosporella Deighton. The two genera are distinguished by.the mode of proliferation of the conidiogenous cell, percurrent and annellidic in Annellospermosporella, sympodial in Spermosporella.
Spermosporella contains two species hyperparasitic on colonies of tropical Meliola spp. (Deighton 1969; Deighton & Pirozynski 1972), and another described from conidiophores of Haplographium Berk. & Broome from Canada (Sutton 1973). The Meliola-inhabiting species are distinguished from a suite of other hyperparasites on Meliolaceae by having hyaline, short, sympodial conidiophores aggregated into sporodochial-like masses, and hyaline conidia with an attenuated, curved apex-features shared by Annellospermosporella meliolinae. Although A. meliolinae is likely to be closely related phylogenetically to the Spermosporella species, it has been placed in a distinct genus because of the significance which continues to be given to small differences in patterns of conidiogenesis in hyphomycete systematics (Sutton & Hennebert 1994).

TYPE SPECIES: Annellospermosporella meliolinae P.R.Johnst.
SPECIMENS EXAMINED: NEW ZEALAND: NORTH ISLAND: NORTHLAND: Russell, on Meliolina novae-zealandiae on Metrosideros excelsa, J. M. Dingley, 16 Sep 1967, PDD 68410. BAY OF PLENTY: Rotorua, Tarawera Falls, on Meliolina novae-zealandiae on Metrosideros excelsa, E. H. C. McKenzie, 11 May 1993, PDD 68409.
DESCRIPTION: Colonies develop amongst Meliolina novae-zealandiae hyphae. The colonies are macroscopically white or pale brown, initially with the appearance of small tufts of hyphae and conidia near the tips of the upright Meliolina hyphae, but with time appearing to cover much of each Meliolina hypha. Hyperparasite hyphae grow within infected host hyphae, emerging at the apex following breakdown of the wall at the tip of the erect Meliolina hyphae. Emergent mycelium 2.5-4.5 µm diam., closely septate, irregularly branched, with pale brown, smooth walls. Conidiogenous cells held singly on the pale brown external mycelium, 10-20 x 3 µm, often irregularly bent, proliferation sympodial, with several denticulate conidiogenous loci. Conidia 20-31 x 3-4.5 µm, straight, fusoid, base truncate, (1-)3-septate, hyaline.

Eriomycopsis species are hyperparasitic hyphomycetes associated with Meliola Fr. and other leaf pathogens. Deighton & Pirozynski (1972) redescribed the genus, treating most of the species described to that time and transferring several to other genera. The genus is characterised by colourless or pale mycelium, polyblastic, denticulate conidiophores, and more or less fusiform, several-septate, hyaline conidia. E. meliolinae is the only species to have been described from Meliolina, and has previously been reported only from Africa (Hughes 1993).

NOTES: There are some differences between the New Zealand material of E. meliolinae and the published descriptions of Hansford (1947) and Deighton & Pirozynski (1972). The mycelium is pale brown rather than hyaline. The arrangement of the conidiogenous cells is distinct from the description of Deighton & Pirozynski (1972), who described and illustrated the conidiogenous cells being held on elongate, several-septate conidiophores. In the New Zealand material the mycelium grows within the hyphae of Meliolina novae-zealandiae, emerging near the tips of the upright hyphal branches. The hyaline conidiogenous cells develop directly on small groups of pale brown cells of irregular shape and size, which develop around the upper part of the host hyphae. Observation at the margins of Eriomycopsis colonies suggests that the hyperparasite initially emerges from the tips of Meliolina hyphae, but that with time its external hyphae may cover a large part of the host hyphae.
The New Zealand material was not associated with the violet soluble dye noted by Hansford (1947). However, Deighton & Pirozynski (1972) noted that there were several distinct hyperparasitic fungi on the E. meliolinae type collection, and it is possible that the dye was associated with another of these fungi. Both of the other Meliolina hyperparasites described in this paper are associated with a deep red or violet dye when mounted in KOH.
Examination of collections of Meliolina novaezealandiae in Herbarium PDD revealed two hyphomycetous hyperparasites in addition to E. meliolinae, but their identity remains unresolved. Both of the unidentified species form bright, white patches amongst the dark Meliolina colonies. Both have hyaline, narrow, thin-walled hyphae which closely envelop the dark and thick-walled Meliolina hyphae. Both species have simple, long-cylindric conidiogenous cells arising directly from the vegetative hyphae, and hyaline, aseptate conidia about 4-6 x 2 µm. In one species (present on PDD 18495, 62067) the conidia are oblong-elliptic with rounded ends, and there is a prominent, flaring collarette at the single, apical conidiogenous locus. The second species (present on PDD 62603) has rhomboid-elliptic conidia with truncate ends, with the conidia often held in short chains at the apex of the conidiogenous cells. The wall of the conidiogenous cell is only slightly thickened at the single, apical conidiogenous locus. Many additional species of hyperparasites would no doubt be revealed by a systematic search for these fungi on the three Meliolina spp. in New Zealand.

SPECIMENS EXAMINED: NEW ZEALAND: NORTH ISLAND: AUCKLAND: North Shore City, Torbay, Stredwick Drive Reserve, on Meliolina leptospermi on Leptospermum scoparium, P. R. Johnston, 14 Sep 1994, PDD 63406. STEWART ISLAND: Mason Bay, near start of track to Freshwater Landing, P. R. Johnston & E. H. C. McKenzie, 23 Nov 1998, PDD 69825.
DESCRIPTION: Colonies comprise a dense mat of closely interwoven, brown-walled hyphae covering the black, hairy Meliolina leptospermi colonies. Malacaria ascomata develop later on the upper surface of the mycelial mat; 0.2 mm diam., globose, dark-brown, with a small, indistinct, round, apical pore. The Meliolina colonies eventually become completely enveloped and apparently replaced by Malacaria. Ascomata 170-200 µm diam. in vertical section, with wall 15 µm thick, comprising mostly angular to short-cylindric 3.5-4.5 µm diam. cells; cell walls slightly thickened, pale brown towards outside of ascoma, hyaline towards inside. Ascomatal wall covered with a thin, incomplete, loose outer layer of tangled, cylindric, 4-5 µm diam. hyphae, their walls thickly encrusted with dark brown material. Winered pigment diffuses into KOH from the ascomatal wall, and possibly also from vegetative hyphae around the ascomata. Lower half of ascoma lined with asci intermixed with persistent, filiform pseudoparaphyses. Asci 95-115 x 9-9.5 µm, subclavate with apex broadly rounded, wall slightly thickened at apex, with a broad, central pore, 8-spored. Ascospores 60-75 x 3 µm, slightly isthmioid, tapering to both ends, 3-5-septate, hyaline, pale brown in mass. Pseudoparaphyses 2 µm diam., mostly unbranched, numerous, persistent.

Pirozynski (1977) discussed a group of ascomycetes hyperparasitic on meholaceous leaf pathogens which have in common dark-walled ascomata with diffusible red or blue piµments, and one- to several-septate, hyaline or slightly piµmented ascospores. Included amongst these was the genus Malacaria, considered by Pirozynski (1977) to possibly be congeneric with Nematothecium Syd. & P.Syd. Rossman (1987), however, retained Malacaria and Nematothecium as distinct. Malacaria is a member of the Tubeufiaceae (Rossman 1987). Nematothecium, having discomycete-like ascomata with a thin, poorly developed wall and sparse pseudoparaphyses, was provisionally placed in the Pseudoperiporiaceae by Eriksson & Hawksworth (1993, as Dimeriaceae) and Hawksworth et al. (1995), but excluded from that family by Barr (1997).
Despite the taxonomic confusion surrounding this group of fungi at the generic and higher levels, many similar species have been described as hyperparasites of Meliolaceae and other biologically and macroscopically similar fungi (Hansford 1946, 1954; Sathe & Vaidya 1976; Rossman 1987). Many of these species are poorly known, and often lack type material (Rossman 1987).
The only species of Malacaria to have been described from Meliolina is M. meliolinae Hansf, reported from Uganda and India (Hughes 1993). Although the type of M. meliolinae was not examined by Rossman (1987), Hansford's (1946) description matches Rossman's concept of Malacaria.

NOTES: The New Zealand material closely matches Hansford's (1946) description of M. meliolinae, and is for now considered to belong in this species. The isthmioid ascospore shape (Cannon 1995) has not previously been noted, but the central constriction of the spores is indistinct, and is easily missed for spores still within the asci. The small differences in ascus and ascospore size (cited as 110-130 x 10-12 M and 70-90 x 3 µm, respectively, by Hansford 1946) are here regarded as probable intraspecific variation. However, Hansford's type material is apparently missing (Rossman 1987), and greater assurance as to the significance of these differences will require additional collections from throughout the range of this fungus. Such collections may show this Meliolina hyperparasite to be genetically distinct in the geographically widespread regions from which it has been reported.

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18 March 2001
22 March 2001
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