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Hausner, G.; Eyjólfsdóttir, G.G.; Reid, J.; Klassen, G.R. 1992: Two additional species of the genus Togninia. Canadian Journal of Botany 70(4): 724-734.

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Hausner, G.; Eyjólfsdóttir, G.G.; Reid, J.; Klassen, G.R. 1992: Two additional species of the genus Togninia. Canadian Journal of Botany 70(4): 724-734.
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CULTURES EXAMINED: NEW ZEALAND: 113 bi, isolated from C. macrocarpa, Compartment 14, Woodhill State Forest, Auckland, collected 25 May 1982; 116c', isolated from P. radiata, Compartment 24, Woodhill State Forest, Auckland, collected 25 May 1982; 89 bi and 105 aiii, both isolated from P. radiata, off Road 41, Whangapoua State Forest, Coromandel, collected 19 May, 1982.
ANAMORPH: Variable, appearing intermediate between an Acremonium and a Phialophora species.
Colonies 38-42 mm in diameter in 21 days at 20°C in darkness on MEA-YE. Colonies grown in darkness gray (5Y 5/1) at the centre, pale yellow to light gray (5Y 8/4 to 5Y 7/2, 2.5Y 7/2) towards the margins, and finally white (10 YR 8/1) at the margins. Colonies appressed, nematogenous to finely plectonematogenous, and consisting of hyphae and hyphal strands bearing short phialides with clear droplets containing conidia at their apices. Colonies grown in alternating light and darkness light gray, light brownish gray to gray, and light olive gray (10YR 7/1, 7/2, 6/2 to 5Y 6/1 and 6/2), often with gray to dark grayish brown (l0YR 5/1 to 4/2) sectors. On aging patches of white, loosely organized phialophores may develop. Cultures grown in the dark for 3 weeks, then transferred to alternating light and darkness for an additional 3 weeks, develop brown hyphal aggregations and brown to black protoperithecia on the surface; the latter develop into mature perithecia in a further 4 to 6 weeks. In reverse, dark grown colonies are gray (5Y 6/1 to 5/1) in the centre, white to pale yellow (5Y 8/2 to 8/3) towards the margin, but become olive gray (5Y 5/2) to dark gray (2.5Y 4/0) on aging. Odour indistinct. Exudate may be present as small clear drops. Hyphae hyaline to brown; smooth to verruculose; 1.5-4.0(5.0) µm in diameter; when funiculose, strands are 5-20 µm in diameter and consist of 1.7-3.0 µm wide hyphae. Chlamydospores not seen. Phialophores micronematous to semimacronematous; hyaline to brown; when brown, darkest towards the base; walls smooth to verrucose; 20-75 x 2.4-2.8 µm including the terminal phialides; each bears 1-2 terminal phialides that are subtended by an axis of 1-3 cells, each of the latter cells sometimes having a lateral adelophialide just below its apical septum (Fig. 3k); or the axis is lacking. Complex, indeterminate, loosely arranged phialophores (Fig. 3l) that are hyaline to pale brown at the base, form in aging cultures. These consist of subulate phialides that have proliferated terminally several times and sometimes give rise to lateral branches (Fig. 3l, arrow). Conidiogenous cells monophialidic or rarely with a lateral locus; hyaline to brown; smooth to verruculose; integrated or discrete. Phialides of three types: (i) adelophialides that are either cylindrical and 2.0-8.5(12.0) x 1.0-2.3 µm, or occasionally widest at the base and tapering to the apex (Fig. 3i); (ii) obclavate to navicular and often curved, narrowing abruptly to a short neck 7.5-15.5 x 2.2-3.0 µm, and tapering to 1.0-1.6 µm at the apex (Fig. 3j); (iii) subulate or slightly inflated at the base, narrowing gradually to a long neck 16-33 x 1.6-2.4 µm, tapering gradually to 1.0-1.6 µm at the apex (Figs. 3k, 3l). All develop 1.0-1.8 µm long indistinct collarettes with periclinal thickenings that may make the apex slightly wider than the neck below. Phialoconidia aggregate at the tips of individual phialides in slimy drops; 1-celled, hyaline, and smooth-walled; most are cylindrical to allantoid and measure 3.5-7.5(9.0) x 1.4-2.4 µm; and a few are oval to short-elliptical and measure 2.4-4.8 x 1.5-2.4(2.8) µm; apex rounded, base indistinctly apiculate on the thinner conidia but rounded on the broader ones (Fig. 3m). Larger conidia that are broadly oblong-elliptical to reniform and (5.0)6.0-10.5 x 2.4-3.2(3.4) µm often formed within the medium; these developed a short adelophialide that produced smaller conidia (microcyclic conidiogenesis) (Fig. 3n). Perithecia developing singly or in clusters; bases globose to broadly oval; black; 175-265 µm in diameter (Fig. 3a); ornamented with short, septate, curved, dark brown, and often fairly thick-walled hyphae (Fig. 3b). Necks 1-2 per perithecium; black but light brown to hyaline at the apices and somewhat curved; surface coarsely verrucose, their apical portions often becoming nodulose upon aging as the neck proliferates terminally; 450-1300 µm long 40-65 µm wide at the base tapering to 35-45 µm at the apex; often widest at the nodes. Asci arising from the ascogenous hyphae that each produce up to at least 15 asci in acropetal succession and continue to elongate in the process; ascogenous hyphae are hyaline, smooth-walled, 1.8-2.5 µm wide, with an irregular, inflated base 2.5-4.5 µm wide (Fig. 3f). The asci appear to be dehiscent from the generative hyphae when mature (Fig. 3g). Asci 8-spored; clavate with nearly truncate apices and sides that are mostly straight but abruptly tapered below to a truncate base; base often appearing lateral as a result of one side of the ascus being sharply curved; 15.5-24 x 4.7-5.5(6.5) µm. The apical complex is 0.7-1.8 µm thick, but its structure is indistinct. Ascospores extruded from the ostiole in slimy masses; 1-celled, hyaline and smooth-walled; elliptical to oblong-eliptical and slightly rounded at each end; 3.8-5.6 x (1.8)2.2-2.6 µm; biguttulate during late stages of development but the oily droplets may be absent from the mature spores. Paraphyses hyaline, thin-walled, and septate; 60-115 x 4.0-5.5 µm wide at the base, tapering to 2.0-2.8 µm at the apex (Fig. 3c). The paraphyses originate from the same pseudoparenchymous tissue as the ascogenous hyphae that produce the asci. Paraphyses dispersed µmongst the young asci, but as the asci mature, they tend to collapse and become inconspicuous.
HABITAT: Cupressus macrocarpa Hartw., Pinus radiata D. Don.
Coloniis in 21 dies 38-42 mm diametro in temperies 20°C crescentibus; in tenebris griseis vel flavis sed marginibus albis; adpressis, nematogenis vel plectonematogenis; inodoris. Hyphis hyalinis vel brunneis, laevibus vel verruculosis; 1.5-5.0 µm diametro. Phialophoris micronematis vel hemimicronematis, hyalinis vel brunneis; parietibus laevibus vel verrucosis, 20-75 x 2.4-2.8 µm. Phialidibus cylindricis vel contractis, obclavatis vel navicularibus aut subulatis; semper collari inconspicuo praeditis; proliferentibus. Conidiis aut cylindricis vel allantoideis, 3.5-7.5(9.0) x 1.4-2.4 µm, aut ovalibus vel ellipsoideis, 2.4-4.8 x 1.5-2.4(2.8) µm. Peritheciis nunc solitariis nunc fasciculatis; basi globosis vel ovalibus, 175-265 µm diametro; pilis fuscis ornatis aut laevibus; collis ad 1300 µm longis, basi 40-65 µm, apice 35-45 µm latis. Ascis octosporis, clavatis vel cylindricis, truncatis apice basique; 15.5-24 x 4.7-5.5(6.5) µm. Ascosporis unicellularibus, hyalinis, laevibus, ellipticis vel oblongo-ellipticis, 3.8-5.6 x(1.8)2.2-2.6 µm. Paraphysibus basi latissimis, productis, septatis, contractis.
In culture characteristics and phialide shape, the anamorph of T. novae-zealandiae resembles some species of the genus Phialophora Medlar except it only produces indistinct collarettes. Phialide collarettes are a key character of Phialophora species. Gams and McGinnis (1983) reintroduced the genus Lecythophora Nannfeldt (Melin and Nannfeldt 1934) for Phialophora species with adelophialides, but the characters cited by them for those adelophialides are not found in T. novae-zealandiae. The anamorph of T. novae-zealandiae is also excluded from Acremonium Link, section Gliomastix, because the colonies that become darkly pigmented are not difficult to cut through. Tough, leathery, difficult to cut colonies are characteristic of the section Gliomastix (Gams 1971, pp. 73 and 242).
Gams and McGinnis (1983) erected Phialemonium W. Gams and McGinnis for species intermediate between the genera Acremonium and Phialophora. In the former, the adelophialide is the typical situation, and discrete, longer phialides are very rare even on aerial hyphae. In the two latter genera, adelophides may be present in some species, but mainly they occur on submerged hyphae. The anamorphic state of T. novaezealandiae also seems to possess characteristics intermediate between Acremonium and Phialophora. However, our fungus differs from any described Phialemonium species in having broader hyphae, possessing adelophialides with indistinct collarettes but distinct periclinal thickenings, and producing a large number of discrete phialides that may proliferate secondarily. Thus, the anamorph of T. novae-zealandiae may represent another intermediate between Acremonium and Phialophora.
Strain 89 bi differed slightly from the other strains. It grew slightly faster, 50 mm in 21 days, did not sector as much, and in darkness the colonies were not as yellowish. It also produced fewer perithecia, but they and their contents were identical to those of the other three strains of this species.
Light exposure usually stimulated perithecial production, with protoperithecia developing from aggregations of brown superficial hyphae in 5- to 7-week-old colonies. The centrum of developing perithecia consists of short, thin-walled, hyaline, 2.5-3.2 µm wide hyphae (Fig. 3d) that elongate apically to form paraphyses (Figs. 3c, 3e) before the asci start to develop. This occurs prior to any evidence of neck development, but only when the necks begin elongating are developing asci found µmongst the paraphyses (Fig. 3e).
In culture, all strains of T. novae-zealandiae were relatively slow growing. On the natural substrates where perithecia were abundant, it was usually associated with other fungi. These observations suggested it might either be parasitic on, or was obtaining some growth factor from, the other fungi. To test this, strain 113 bi was grown in paired cultures with an isolate of Sclerotium hydrophilum Sacc., Epicoccum nigrum Link, a Cladosporium sp., and a sterile demateaceous fungus isolated from bark beetle galleries. In all cases, perithecia formed in such pairings had longer necks and were slightly more robust than those formed in pure cultures, and T. novae-zealandiae overgrew the other fungus. No evidence of active parasitism was noted, but stimulation was certainly apparent.
HOLOTYPE: NEW ZEALAND: Dried culture of isolate 113 bi, from C. macrocarpa, Compartment 14, Woodhill State Forest, Auckland, collected by J. Reid, 25 May 1982, WIN.

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17 November 2003
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