ARGENTINA: CHUBUT: Parque Nacional Los Alerces, on N. dombeyi, R. Menéndez, I. J. Gamundí & A. J. Amos, 3 Sep 1996, BCRU 1551. NEUQUEN: Parque Nacional Nahuel Huapi, International Milestone, near Road 215, on N. dombeyi, M. I. Messuti, 31 Mar 1993, BCRU 1754; Pla. Quetrihué, on N. dombeyi, I. J. Gamundí & C. Brion, 1 Jun 1997, BCRU 1767; Parque Nacional Lanín, A. Queñil-Hue, on N. dombeyi, I. J. Gamundí, 10 Apr 1992, BCRU 1757; Hua-Hum, on N. dombeyi, I. J. Gamundí, 6 Apr 1992, BCRU 1758. RIO NEGRO: Parque Nacional Nahuel Huapí, Llao-Llao, Parque Municipal near Soria Moria, on N. dombeyi, I. J. Gamundí, A. L. Giaiotti & M. I. Messuti, 28 Apr 1993, BCRU 1753; Path from Pt. Blest to Los Cántaros, on N. dombeyi, P. R. Johnston SA193, I. J. Gamundí & C. Brion, 2 Apr 1996, BCRU 1753, PDD 69811; L. Nahuel Huapi, Brazo Tristeza, on N. dombeyi, I. J. Gamundí, 3 May 1994, BCRU 1750; Lago Mascardi, on N. dombeyi, C. Pujals & I. J. Gamundí, 24 Apr 1958, BAFC 20345; Llao- Llao, on N. dombeyi, C. Pujals & I. J. Gamundí, 27 Apr 1958, BAFC 20304. TIERRA DEL FUEGO: Ushuaia, Glaciar Martial Valley, near base of ski lift, on N. betuloides, P. R. Johnston SA3 & L. Lorenzo, 16 Mar 1996, BCRU 1848, PDD 69810; Ushuaia, Glaciar Martial Valley, ski trail below Tea House, on N. betuloides, P. R. Johnston SA119 & L. Lorenzo, 24 Mar 1996, BCRU 1847, PDD 68405; Parque Nacional Lapataia, Bahía Ensenada, on N. betuloides, P. R. Johnston SA154 & L. Lorenzo, 27 Mar 1996, BCRU 1849, PDD 69807; Bahía Ensenada, Punto Panorámico, I. J. Gamundí, 16 Mar 1975, BCRU 1267; Bahía Ensenada, I. J. Gamundí, 17 Feb 1972, BCRU 1183; Lapataia, peat bog, on N. betuloides, E. Horak, 19 Feb 1974, BCRU 1185; L. Escondido, road Ushuaia to L. Fagnano, P. R. Johnston SA77 & L. Lorenzo, 22 Mar 1996, BCRU 1846, PDD 69809; Estancia Moat, on N. betuloides, I. J. Gamundí, 18 Mar 1975, BCRU 1186; Isla de los Estados, on N. betuloides, I. J. Gamundí, 28 Nov 1967, BCRU 1268; Bahía Ensenada, on N. betuloides, I. J. Gamundí, 17 Feb 1972, BCRU 1183; Bahía Ensenada, on N. betuloides, I. J. Gamundí, 16 Mar 1975, BCRU 1267; Road Ushuaia-Moat, Estancia Moat, on N. betuloides, P. R. Johnston SA134 & L. Lorenzo, 25 Mar 1996, BCRU1850, PDD 69808; Tierra Mayor, on N. betuloides, I. J. Gamundí & P. Medina, BCRU 1184.

CHILE: LOS LAGOS: Prov. Palena, L. Yelcho, track to Glaciar Yelcho, on N. dombeyi, P. R. Johnston SA201 & M. Rajchenberg, 5 Apr 1996, PDD 68406; Prov. Osorno, Parque Nacional Puyehue, road to Antillanca, 1 km up from Lag. El Toro, on N. dombeyi, P. R. Johnston SA245 & I. J. Gamundí, 9 Apr 1996, PDD 68403, BCRU 1998.

Apothecia developing on fallen leaves, growing on both sides and margin of leaves, leaves darkened but not showing definite stromatised areas; erumpent through host cuticle, arising from small, dark stroma; apothecia 0.6-2 mm diam., stipitate, fleshy; disc plane or convex, grey-yellowish ("avellaneus" to "sepia"); receptacle dark brown, receptacle and margin with fuligineous setae; stipe 0.5-1 x 0.18-0.27 mm, cylindrical, base bulbous, blackish. Paraphyses 1.5-2 µm diam., slightly swollen near apex. Asci 117-147 x 9-11 µm, cylindric-clavate, tapering to base, wall thickened at apex, with strongly amyloid pore, 8-spored. Ascospores 16-17.5 x 3.8-5.8 µm, fusoid, asymmetrical, containing 1-4 guttules, hyaline to slightly yellowish; 1-2-seriate. Ectal excipulum 3-layered; outer layer one cell wide, forming a radial network over receptacle, more or less parallel on stipe, comprising hyphae 2.5-5.5 µm diam. with walls brown, thickly encrusted; central layer 18-25 µm thick, comprising textura porrecta of hyphae 2.5-4.5 µm diam. with walls hyaline, gelatinous; inner layer 18-23 µm thick, comprising textura porrecta of hyphae 2.5-4.5 µm diam. with walls pale brown, irregularly encrusted, nongelatinous. Medullary excipulum comprising textura intricata of hyphae 2.5-4.5 µm diam. with walls hyaline, nongelatinous. Setae arising from the central layer of ectal excipulum on receptacle, from outer layer of ectal excipulum on stipe, 150-520 x 5.8-11 µm, swollen near the base, tapering to base, base simple on receptacle, L- or T-shaped on stipe, wall smooth, dark brown, opaque, often paler near subacute apex, pluriseptate.

APPEARANCE IN CULTURE: OA: 18-80 mm diam.; aerial mycelium cottony, forming patchy tufts, white to grey; agar surface orange-brown to red-brown; in reverse with patches of orange brown pigmentation. PDA: 30-35 mm diam.; margin uneven, surface of colony raised and deeply convoluted; aerial mycelium low, felted to cottony, tufted, variable in pigmentation, white, pale grey, bright yellow, or pinkish; in reverse very dark orange-brown; deep yellow pigment diffusing into agar across plate. MEA-M: 20 mm diam.; aerial mycelium sparse, whitish; surface of agar deep orange-brown; orange-brown pigments diffusing into agar across plate (BCRU 1850 with less well developed pigment, faster growth, and quite dense tufted to felted mycelium with irregular patches of white, pale grey, and bright yellow pigments). MEA-D: similar to MEA-M (BCRU 1847 with faster growth, thin colonies with little aerial mycelium and irregular patches of purplish-grey pigment).

ETYMOLOGY: the specific epithet refers to the southern South American distribution of this species, along the Andes.

T. andina is the most common Torrendiella species in the Andean-Patagonian forest of South America. It is characterised by the subcuticular stroma from which the apothecia arise, the long stipe, and very dark receptacle. The dark appearance is due to the well developed outer excipular layer. This layer, continuous and one cell thick in the stipe, forms a brown net over the receptacle. Small, immature apothecia are darker than those that are mature, and it is possible that as the hymenium expands with maturity, the outer excipular layer becomes disrupted and the hyphae become separated, so forming the characteristic net.

Although T. andina is found on more than one host species, it is interesting to note that N. betuloides and N. dombeyi are closely related, and are both evergreen with similar, coriaceous leaves. N. dombeyi grows in northern Patagonia on the eastern slopes of the Andes from about 38^o 40' S to 43^o 40' S and is a montane species; N. betuloides reaches sea level in Tierra del Fuego, growing from about 48° S to 56° S.

Note that the collections BAFC 20345 and 20304 were recorded erroneously by Gamundí (1962) as Zoellneria eucalypti.

Apothecia developing on both sides of fallen leaves, or on dead leaves still attached to fallen branches; erumpent through host leaf epidermis, arising from small patch of compact, hyaline textura intricata at base of stipe; apothecia 0.25-0.4 mm diam., sessile to short stipitate, fleshy; disc plane, grey; receptacle dark grey, colour more intense at base, with scattered, short, black, stiff setae; stipe short, broad, often paler than receptacle. Paraphyses 1.5-2 µm diam., slightly swollen to 3-4 µm diam. at rounded apex, branched, extending about 5 µm beyond asci. Asci (100-)115-140(-150) x (10-)12.5-14.5 µm, cylindric-clavate, apex rounded to subtruncate, wall slightly thickened at apex, with indistinct, diffuse amyloid reaction in outer part of wall, 8-spored. Ascospores (15-)17-20 x 6-7.5 (-8.5) µm, elliptic-fusoid, slightly wider in upper half, flattened on one side, often slightly curved, more or less acute to the often slightly beaked apex, rounded to base, hyaline. Ectal excipulum varies in appearance between apothecia; in some apothecia the layer of gelatinous tissue characteristic of Torrendiella is well developed, while in others it is very poorly developed. In "typical" apothecia outer layer poorly differentiated, comprising meandering hyphae 2 µm diam. with walls finely encrusted and pale brown, forming sinuate pattern across surface of receptacle; central layer up to 40 µm thick, comprising hyphae 2-2.5 µm diam. with walls thin, very pale brown and finely encrusted, widely spaced within hyaline gelatinous matrix; inner layer 10-15 µm thick comprising textura porrecta of cylindric to brick-shaped cells 3-6 µm diam. with walls brown, nongelatinous; at base of stipe, gelatinous tissue less well organised, forming textura intricata of hyphae with walls hyaline, thick, gelatinous. In "atypical" apothecia, gelatinous nature of tissue at sides of excipulum is lost, with the pale brown, finely encrusted hyphae of central layer forming a complete, 1-3 cell wide layer across surface of receptacle; inner layer remains same as in "typical" apothecia, as does gelatinous tissue at base of stipe. Setae arising from central layer of ectal excipulum, 100-130 x 5-7.5 µm, often slightly curved, inflated near the base, tapering to simple base, and gradually to rounded apex, walls smooth, brown, slightly paler near apex, pluriseptate.

ETYMOLOGY: The specific epithet refers to the short setae characteristic of this species.

T. brevisetosa is characterised by its small, dark apothecia, relatively short, somewhat curved setae, and large ascospores. T. dingleyae has similar small, dark apothecia, but it has much longer setae, and differs in ascospore size and excipular structure.

T. brevisetosa is variable with respect to development of gelatinous tissue within the excipulum. This difference appears to relate in part to maturity and aging of apothecia; all immature apothecia (ascospores visible within asci but not fully differentiated) that have been sectioned have a well-developed gelatinous excipulum, most mature apothecia (ascospores fully differentiated within asci) lack the gelatinous layer. However, this is not always the case in the genus, other species with poorly developed gelatinous layers (e.g., T. cannibalensis) showing no variation in mature versus immature apothecia.

The type collection is from leaves that contain a mixture of T. brevisetosa and T. cannibalensis apothecia. Although the two species sometimes occur adjacent on a single leaf, in most cases the T. brevisetosa apothecia are on the upper surface of the leaves, the T. cannibalensis apothecia on the lower surface. T. cannibalensis has narrower ascospores, and an ascus pore with a more intense amyloid reaction.

A collection from fallen leaves of Nothofagus cunninghamii from Australia (Victoria, Melba Gully, P. R. Johnston AU96-2 & B. Fuhrer, 17 May 1996, MEL) is macroscopically similar in having small, dark apothecia with a pale stipe and short setae, a central excipular layer in which the hyphae are held apart in a hyaline gelatinous matrix, and asci with a similar, diffuse amyloid reaction at the apex (Fig. 5). The Australian species differs in having slightly longer asci (140-160 µm), and slightly shorter and broader ascospores (14.5-16.5 x 6.5-8.5 µm) which are ovate and symmetrical in shape. Known from a single, mostly immature collection, the species from Australia will not be formally described here.

Apothecia developing on both sides of fallen leaves; erumpent through host epidermis, arising from small patch of compact tissue comprising angular cells with hyaline walls, intermixed in part with host hypodermal cells; apothecia 0.3-0.8 mm diam., short-stipitate, fleshy; disc plane to slightly convex, translucent, greyish ("beige"), drying darker ("isabellinus"); receptacle slightly darker than hymenium, with reddish tint, with dark, stiff, setae; stipe short, obconical, 0.5 mm long, 0.2-0.25 mm diam., concolorous with receptacle. Paraphyses 1.5-3.5 µm diam., undifferentiated or slightly and irregularly swollen near rounded apex, sometimes 1-branched, some containing yellowish guttules in upper part. Asci (75-) 85-110 (-130) x (7.5-) 9.5-11.5 µm, cylindric to subclavate, tapering slightly to rounded to subtruncate apex, wall slightly thickened at apex, pore with intense amyloid reaction, 8-spored. Ascospores (14.5-) 17-20 (-23) x 4-5 µm, elliptic, asymmetrical, flattened one side, slightly curved, slightly wider in upper half, tapering to more or less acute apex, more rounded toward base, hyaline; overlapping 1-2-seriate. Ectal excipulum 3-layered; outer layer poorly developed, comprising loose network of hyphae 2-3.5 µm diam. with walls hyaline to pale brown, with often numerous, short, irregular divarications, apical cells of hyphae often with dense contents; central layer up to 40 µm thick, confined to stipe and base of receptacle, on receptacle comprising regular rows of cylindric cells 2.5-4 µm diam. with walls hyaline, thickened, gelatinous, this layer becoming partly lost and less well organised in older apothecia, where it is confined to base of stipe, and comprises angular to subglobose cells with walls hyaline, thickened, gelatinous; inner layer 10-30 µm thick, comprising rows of long-cylindric cells 4-8 µm diam. with walls brown, thickly encrusted, nongelatinous. Setae arise from central layer of ectal excipulum, (120-) 150-180 (-220) x 5.5-7.5 µm, swollen near the base, tapering to base, tapering slightly to broadly rounded apex, walls smooth, brown but not opaque (septa visible under microscope) in lower part, upper 3-4 cells pale brown to hyaline.

APPEARANCE IN CULTURE: OA: 80 mm diam.; aerial mycelium sparse, in small scattered tufts somewhat concentric in arrangement, pale grey; agar surface orange-brown; in reverse deep orange-brown. PDA: 70 mm diam.; aerial mycelium dense, finely tufted, whitish; in reverse deep reddish-brown. MEA-M: 50 mm diam.; aerial mycelium low, dense, crusty, whitish; agar surface orange brown; in reverse deep orange brown; yellowish pigment diffusing into agar across plate. MEA-D: 55 mm diam.; thin colony with sparse mycelium, pale brownish pigment near centre.

T. cannibalensis is macroscopically distinct amongst the Nothofagus-inhabiting species in possessing short-stipitate apothecia with a very pale receptacle, contrasting with dark setae. T. eucalypti has a similar pale apothecium, but they are more distinctly stipitate, and the setae are longer and less numerous. Excipular structure and appearance in culture also differ between T. eucalypti and T. cannibalensis.

Two other Torrendiella species on Nothofagus in New Zealand have small, macroscopically similar apothecia, T. brevisetosa and T. dingleyae. However, the ascospores of T. brevisetosa are wider (6-7.5 µm), while the ascospores of T. dingleyae are symmetrical. T. cannibalensis is also distinguished by its characteristic excipular structure; the outer layer on the receptacle comprises hyphae with irregularly divaricating cells, while the central gelatinous layer is confined mostly to the base of the apothecium. The excipulum comprises mostly cylindric, brown-walled, nongelatinous cells, characteristic of the inner layer of the ectal excipulum of Torrendiella.

Apothecia developing on both sides of fallen leaves or on dead leaves remaining attached to fallen twigs and branches; erumpent through host tissue, arising from small patch of stromatic tissue at base of apothecium; apothecia 0.35-0.65 mm diam., short-stipitate, fleshy; disc plane, sepia grey or light greyish sepia; receptacle and stipe concolorous with disc (darker in some collections), with numerous, black, stiff setae; stipe robust, short, cylindric, often darkened near base. Paraphyses 1.5-2 µm diam., more or less undifferentiated at rounded apex, about same length as asci. Asci 105-135 (-145) x 7.5-9 µm, cylindric, tapering slightly to broadly subtruncate apex, wall thickened at apex with prominent amyloid pore, 8-spored. Ascospores (11.5-) 12.5-14.5 (-15.5) x (4.5-) 5.5-6.5 (-7) µm, broad ellipsoid-fusoid, tapering uniformly to each end, hyaline; uniseriate. Ectal excipulum 3-layered; outer layer a single cell wide network of radially disposed, long-celled hyphae 2.5-5 µm diam. with wall encrusted, dark brown, end cell often slightly swollen; central layer comprising textura porrecta of long-cylindric cells with lumen 2.5-3 µm diam. on sides of receptacle, 5-6.5 µm diam. near base, wall hyaline, thick, gelatinous; inner layer comprising textura porrecta of long-cylindric cells 4-8 µm diam. with wall brown, thin, nongelatinous. Setae arising from central layer of ectal excipulum, (300-) 400-500 (-600) x 12-15 (-20) µm, slightly swollen near the base, tapering to simple base, tapering gradually to more or less acute apex, walls smooth, dark brown to black, slightly paler near apex, pluriseptate (but septa visible only in pale upper portion of setae); setae on stipe shorter and thinner than those on receptacle, about 190-230 µm long, tapering to either simple base, or irregularly swollen at base, those with swollen base arising from outer layer of ectal excipulum.

APPEARANCE IN CULTURE: OA: 80 mm diam.; aerial mycelium dense, grey, forming irregular patches, lacking from most of colony; agar surface patchy with black, dark brown, and red brown pigmentation. PDA: 35 mm diam., margin slightly uneven; colony surface near centre raised and deeply convoluted; aerial mycelium low, dense, felted to cottony, whitish to pale grey; in reverse very dark reddish brown. MEA-M: 75 mm diam.; aerial mycelium dense felted to cottony, pale grey to pinkish; in reverse pale reddish brown, with irregular dark brown patches. MEA-D: aerial mycelium sparse; agar with brown pigmentation, more intense toward centre of colony, with small, scattered dark-brown flecks.

T. grisea, known only from Nothofagus dombeyi, is characterised by the pale brown setae that are swollen and hyaline at the apex. Although unusual for Torrendiella, at least one other species, an undescribed species from Epacridaceae from Australia and New Zealand, has the same kind of setae (P.R. Johnston, unpubl. data). The setae of these fungi are very like those of Zoellneria rosarum, the type of Zoellneria; however that genus differs from Torrendiella in other respects (see Introduction). The 3-layered nature of the ectal excipulum is poorly developed in T. grisea.

Note that the collections BAFC 20267a and 20347 were recorded erroneously by Gamundí (1962) as Zoellneria eucalypti.

AUSTRALIA: NEW SOUTH WALES: Northern Tablelands, Gloucester Tops, creek below Darby Munro Hut, on N. moorei wood, R. Filson, 26 Oct 1963, MEL 257720. TASMANIA: Geeveston, Hermons Rd, E side, 1.1 Km N of St Johns Rd, on wood, J. Weller, 20 Nov 1995, MEL 2060918; Geeveston, 350m W of Hermons Rd, 150 N Hermons Spur 3 Rd, on wood, T. May 1124, 8 Apr 1984, MEL 2060917; Geeveston, Rutherfords Rd, SE side, N of Johns Creek Crossing, 1 Km S of Johns Rd, on wood, J. Weller, 21 Nov 1995, MEL 2060919. VICTORIA: Melba Gully, near Lavers Hill, on unidentified log, G. Beaton 47 (E0405), 8 Dec 1962, MELU 99.1.1 - holotype; Otway Range, Melba Gully, on fallen wood, G. Beaton, 25 Feb 1963, MEL 2032728.

NEW ZEALAND: FIORDLAND: Fiordland National Park, Hollyford Rd., Lake Marian track, on Nothofagus sp. wood, P. R. Johnston D776, 3 Mar 1992, PDD 59983. TAUPO: Kaimanawa Forest Park, Clements Rd, on N. menziesii wood, P. R. Johnston D299, 22 Nov 1988, PDD 54837; Kaimanawa Forest Park, Clements Rd, on Nothofagus sp. bark, P. R. Johnston D759 & R. E. Halling, 23 Feb 1992, PDD 60516. WESTLAND: vic. Haast, Lake Ellery, on Nothofagus sp. wood, P. R. Johnston D655, 21 Mar 1991, PDD 58589; vic. Haast, Lake Paringa, on Nothofagus sp. bark and wood, P. R. Johnston D672, 22 Mar 1991, PDD 58572.

Apothecia developing on decorticated wood and bark, surface of colonised wood sometimes blackened; apothecia up to 5 mm diam., stipitate, fleshy, tough; disc plane, initially pale grey, deep brownish-yellow with age; receptacle pale yellow-brown to almost black, receptacle covered with numerous, short, fuliginous, somewhat flexuous setae; stipe tapering toward base, dark, often densely covered with pale, flexuous setae. Paraphyses 2-3 µm diam., more or less undifferentiated to apex, often containing dense yellowish material, this released as yellow pigment into KOH. Asci 90-130 x 6.5-7.5 µm, narrow-cylindric, tapering to rounded apex, wall thickened at apex, with elongate, often weakly amyloid pore, 8-spored, with long basal stalk. Ascospores (10-) 12.5-16 (-17) x 3.5-4.5 µm, elliptic-fusoid, inequilateral, slightly flattened one side, slightly wider in upper half, ends rounded, sometimes becoming 1-septate, overlapping uniseriate, often producing small, simple ascoconidia from each end of spore. Ectal excipulum 3-layered; outer layer thin, comprising long cylindric cells, nongelatinous, some cells hyaline, 2-3 µm diam., commonly with short divarications, superimposed by network of brown hyphae 4-5 µm diam. with walls encrusted with dark brown material; central layer comprising tangled, cylindric cells 4-6 µm diam. with walls hyaline, somewhat thickened, agglutinated; inner layer up to 45 µm thick, comprising hyphae 3-4 µm diam. with walls thin, finely encrusted, pale brown, non-gelatinous. Setae arising mostly from dark-walled cells of outer layer of excipulum, but from central layer of excipulum at margin of disc, (100-) 180-400 x 4-7 µm, tapering gradually to rounded apex from base, T- or L-shaped at base, walls smooth, brown towards base, becoming paler toward apex, with uppermost 2-3 cells more or less hyaline, pluriseptate.

APPEARANCE IN CULTURE: OA: 80 mm diam.; aerial mycelium sparse; agar surface variable in pigmentation with black or whitish patches, or pigmentation may be more or less lacking completely, or entire colony may be black. PDA: 70 mm diam.; aerial mycelium lacking; agar surface with irregular patches of dark grey-brown pigment. MEA-M: 80 mm diam.; aerial mycelium low, tufted, arranged in narrow concentric bands, mostly dark grey but with scattered patches with brighter orange-brown colours; in reverse dark grey-brown to grey-purplish. MEA-D: 70 mm diam., aerial mycelium sparse, dark; agar uniformly pigmented dark grey-brown.

T. madsenii is characterised by large apothecia with a broad stipe, narrow, often flexuous setae that commonly have a T-shaped base, and ascospores that often produce ascoconidia. In addition, this is the only Nothofagus-inhabiting species that grows on wood rather than leaves. Previously reported only from Australia (Spooner 1987), the geographical range of this species is widened to New Zealand and southern South America. The stromatic development on the host surface noted by Spooner (1987) for the Australian collections was not present on the collections from New Zealand. Note that the Argentinian collection Singer M3005 was recorded erroneously by Gamundí (1962) as Zoellneria eucalypti.

All collections of this species from New Zealand and South America have been on wood of Nothofagus. Although the host preference of the Australian collections remains uncertain, all the collecting sites reported by Beaton & Weste (1977) and Spooner (1987) have Nothofagus present in patches (T. May, Melbourne Botanic Gardens, pers. comm.), as do those of the additional more recent collections examined from MEL. Torrendiella collections in MEL from wood from sites where Nothofagus is lacking all represent T. clelandii or the T. clelandii-like fungus discussed by Spooner (1987). T. clelandii and T. cf. clelandii may be restricted to Eucalyptus. Apart from the difference in preferred substrate, they can be distinguished from T. madsenii by their larger asci and ascospores, and by having setae unbranched at the base.

Most collections of T. madsenii have been made during summer months, unusual for Torrendiella in the Southern Hemisphere.